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A  Study  of  Some  Factors 
Influencing  Fertility  and  Sterility 

in  the  Bull 


^^^X 


^^  - 


BY 

HERBERT  L.  OILMAN 


Reprinted  from  the 
Annual  Report  of  the  New  York  State  Veterinary  College 

1921-22 


...     •      •  ».    '      •    • 


A  Study  of  Some  Factors 
Influencing  Fertility  and  Sterility 

in  the  Bull 


A  THESIS 

Presented  to  the  Faculty  of  the  Graduate  School 

of  Cornell  University  for  the  Degree  of 

Doctor  of  Philosophy 


BY 

HERBERT  LESTER  OILMAN,  D.  V.  M.,  M.  S. 

ii 


Reprinted  from  the 
Annual  Report  of  the  New  York  State  Veterinary  College 


ALBANY 

J.  B.  LYON  COMPANY,  PRINTERS 

1922 


/.  : : .;  ••; .«. •**. 


rv«»M '« Mftit 


'1 


A  STUDY  OF  SOME  FACTORS  INFLUENCING  FERTILITY 
AND  STERILITY  IN  THE  BULL 

HERBERT  L.  OILMAN" 
Veterinary  Experiment  Station,  Cornell  University 

Normal  reproduction  is  the  fundamental  foundation  upon  which 
the  entire  cattle  industry  rests.  For  this  reason,  any  factor 
capable  of  interfering  with  it  is  a  detriment  to  the  industiy,  and 
a  matter  of  prime  importance  to  the  breeder  and  the  veterinarian. 
With  the  relative  increase  in  number  and  value  of  cattle,  an.l  the 
fact  that  the  profession  is  depending  more  and  more  on  this  in- 
dustry for  a  livelihood,  these  problems  are  assuming  greatei-  im- 
portance. The  part  plaj^ed  by  the  bull  has  been  emphasized 
entirely  too  little,  with  the  result  that,  as  in  human  medicine, 
many  fail  to  appreciate  the  effects  of  sterility  or  lowered  fertility 
in  the  male.  The  part  played  by  the  sire  in  the  spread  of  genital 
infections,  though  discussed  frequently,  has  received  little  sys- 
tematic investigation. 

The  bull  miLst  be  regarded  as  at  least  half  the  herd,  not  only 
from  the  standpoint  of  the  characters  he  imprints  upon  his  prog- 
eny, but  because  of  his  relation  to  the  reproductive  efficiency  in 
the  herd.  It  seems  quite  probable  that  he  does  disseminate  during 
copulation,  infection  associated  \\i\h.  the  genital  organs,  with  tlie 
result  that  the  bull  is  a  very  important  factor  in  a  study  of  the 
subject.  Too  frequently,  his  ability  to  copulate  in  an  apparently 
normal  manner,  is  taken  as  a  standard  of  fertility.  Gross  changes 
in  his  genitalia,  or  the  absence  of  spermatozoa  from  the  semen 
are  given  due  consideration,  while  other  more  ol^seure  al)nor- 
malities  are  not  looked  for  nor  regarded  in  their  proper  light. 
Neither  fertility  nor  sterility  are  always  absolute,  but  the  terms 
should  be  used  relatively  inasmuch  as  we  may  have  all  degrees  of 
infertility  or  impotency.  All  Too  frequently  we  foi'get  the  many 
delicate  and  intricate  mechanisms  involved  in  ihe  reproductive 
process,  with  the  result  that  many  phases  of  the  problem  are  neg- 
lected or  disregarded.  The  genital  organs  woi'k  as  a  unit,  each 
part  of  which  must  function  in  perfect  accord  with  the  others  to 
the  end  that  full  fertility  may  result.  The  physiological  factors 
involved  in  the  formation  of  the  semen  are  too  little  understood, 
or  at  best,  our  knowledge  regarding  them  is  more  or  less  hazy. 

The  purposes  of  the  present  work  have  been:  (1)  to  summarize 
the  work  so  far  done  on  the  subject.  (2)  to  review  brietly  the 
known  facts  throwing  light  on  the  anatomy  and  physiology  of  the 
male  genital  organs,  (3)  to  carry  out  systematic  studies  upon  the 
pathology  and  bacteriology  of  the  genital  tract  of  the  bull,  and 
(4)  to  ascertain  if  possible  whether  the  ])ull  is  a  disseminator  of 
those  infection!^-  which  interfere  with  reproduction  in  the  female. 

f3l 


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-•-.  •'    .' .  ♦.'    ::  ..:.••'.  4/ 

Tlie  work  has  been  carried  on  for  the  most  part  from  the  point 
of  view  of  a  laboratory  man  cooperating  with  clinicians.  No 
attempt  is  made  in  this  paper  to  give  detailed  clinical  data,  methods 
for  physical  examinations,  etc.  There  are  included  many  state- 
ments "and  some  data  given  in  a  preliminary  article  on  the  sub- 
ject. While  the  subject  is  broad  in  its  scope,  in  fact  too  broad  for 
great  detail,  it  is  hoped  that  a  start  has  been  made  toward  future 
and  more  detailed  investigations. 

History 
References  to,  and  investigations  relating  to,  the  part  played  by 
the  bull  in  the  process  of  reproduction  in  the  herd,  and  in  the 
spread  of  genital  infections,  have  been  limited  largely  to  those 
phenomena  caused  by  Bad.  ahortum.  Bang  (1)  originally  called 
attention  to  the  possibility  of  the  male  transmitting  the  organism 
discovered  by  him,  but  he  reached  no  definite  conclusion  on  the 
subject.  James  Law  (2)  writing  on  contagious  abortion  in  cows, 
early  suspected  this  possibility  when  stating  under  "  casual  infec- 
tions," that — "  In  a  case  which  came  under  the  observation  of  the 
Avriter  recently,  a  family  cow,  kept  in  a  barn  where  no  abortion 
had  previously  occurred,  was  taken  for  service  to  a  bull  in  a  herd 
w^here  abortion  was  prevailing,  and  though  she  Avas  only  present 
at  the  latter  place  for  a  few  minutes,  she  aborted  in  the  sixth 
month."  Jansen,  as  quoted  by  Sand,  reports  the  case  of  a  cow 
from  an  aborting  herd  having  been  taken  into  a  herd  that  had 
been  previously  quite  free  from  the  disease.  Soon  after  her  arrival 
she  aborted,  and  later  cow  after  cow  of  the  original  herd  aborted. 
The  owner  kept  the  matter  a  secret,  and  sent  his  cow  to  a  neigh- 
bor's bull  for  service,  with  the  result  that  for  two  years  abortion 
prevailed  among  cows  served  by  this  bull.  McFadyean  and  Stock- 
man (3)  later,  in  experimental  work,  attempted  but  failed  to  in- 
fect cows  by  using  a  soiled  bull  for  service.  Hadley  and  Lothe 
(4)  state:  "A  large  number  of  stockmen  hold  that  the  bull  is  an 
important  factor  in  the  transmission  of  contagious  abortion  in 
herds.  A  smaller  number  believe  that  the  bull  merely  acts  as  a 
passive  carrier  of  the  abortion  disease  and  is  not  actively  concerned 
in  the  transmission."  In  a  subsequent  bulletin,  Hadley  (5)  re- 
marks :     ' '  The   abortion   organisms  may  enter   the  body    

during  sexual  intercourse."  In  an  experiment  carried  on  by  the 
same  author  and  co-workers,  abortion-free  virgin  heifers  were  mated 
to  abortion-infected  bulls,  infection  being  evidenced  by  positive 
reactions  to  the  complement  fixation  and  agglutination  tests.  His 
results  indicate,  he  believes,  "  that  the  bull  is  not  so  important  a 
factor  in  transmitting  abortion  as  many  believe."  The  con- 
clusions are:  "Bulls  may  become  infected  with  abortion  bacilli. 
Bulls  that  reacted  to  the  blood  tests  were  incapable  of  dissemi- 
nating the  abortion  disease  to  the  abortion-free  heifers  with  which 
they  were  mated.  Bulls  appear  to  possess  a  sexual  or  individual 
immunitv  to  abortion  infection  that  renders  them  less  susceptible 


than  cows  and  induces  a  milder  form  of  the  disease.  The  resist- 
ance appears  to  be  due  to  certain  anatomic  and  physiologic  differ- 
ences in  their  Sexual  organs  which  make  them  less  favorable  places 
for  the  growth  of  the  abortion  germs  than  those  of  the  opposite 


sex." 


Buck,  Creech,  and  Ladson  (6)  applied  the  agglutination  test  to 
325  mature  bulls,  of  which  288  were  negative  and  37  positive. 
Bacillus  abortus  Avas  isolated  from  five  animals,  of  which  three 
showed  marked  lesions,  two  in  the  seminal  vesicles,  and  one  in  the 
left  testicle.  They  conclude:  "  B.  abortus  may  involve  organs  of 
the  generative  apparatus  of  bulls,  producing  chronic  inflammatory 
changes.  Of  the  generative  organs,  the  seminal  vesicles  appear  to 
furnish  the  most  favorable  site  for  the  lodgement  and  propagation 
of  abortion  infection." 

Schroeder  and  Cotton  (7)  cite  the  case  of  a  bull  which  reacted 
to  the  abortion  test  and,  on  post  mortem,  Bact.  dbortum  was 
isolated  from  an  abscess  of  one  epididymis.  They  state:  "Our 
attempts  to  produce  a  similar  case  of  infection  artificially  failed, 
and,  in  agreement  with  the  difficulties  many  investigators  have 
had  to  obtain  incriminating  CAddence  against  bulls,  we  have  thus 
far  failed  to  infect  bulls  in  any  way  that  justifies  the  assumption 
that  they  are  important  factors  in  the  dissemination  of  abortion 
disease."  Further,  they  conclude:  "  Regarding  the  dissemination 
of  abortion  disease  by  bulls,  we  may  say,  however,  that  it  would 
be  foolhardy  in  the  dim  light  of  our  present  knowledge  to  take 
liberties  with  reacting  bulls,  or  bulls  from  infected  herds,  or 
promiscuously  used  bulls." 

Cotton  (8)"^ failed  to  demonstrate  the  presence  of  abortion  bacilli 
in  the  genital  organs  of  the  bull  used  to  serve  aborting  cows,  or  in 
the  testicles  of  two  bull  calves,  one  of  which  had  been  fed  and  the 
other  injected  M'ith  the  cultures  of  the  abortion  bacillus.  He  con- 
cludes that  the  bull  does  not  harbor  the  organisms  in  the  testicles. 
Carpenter  (9)  injected  both  streptococci  and  Bact.  abort um  into 
the  scrotal  sacs  of  young  calves,  and  intravenously  in  others.  In 
no  case  was  he  able  to  recover  the  organisms  from  any  part  of  the 
genital  canal,  except  for  a  streptococcus  in  one  instance.  Tiettger 
and  AYhite  (10)  were  unable  to  obtain  evidence  of  the  presence  of 
Bact.  abortum  in  three  bulls  slaughtered  after  repeated  reactions 
to  the  complement  fixation  and  agglutination  tests.  The  three 
bulls  had  been  under  observation  for  three  years,  with  no  conclu- 
sive evidence  to  indicate  that  they  were  a  source  of  danger  to  the 
herds  in  which  they  were  a  part.  They  believe  that  the  bull  trans- 
mits the  infection  as  a  passive  carrier. 

Attempts  at  artificial  inoculation  by  natural  channels  have 
failed,  with  the  possible  exception  of  McFadyean,  Sheather,  and 
]\rinett  (11)  who  were  able  to  infect  the  bull  by  the  prepuce  in  two 
cases  and  by  the  mouth  in  one  case.  The  results,  however,  are  by 
no  means  conclusive.  They  conclude,  nevertheless,  that  catth;  of 
any  age  of  either  sex  may  be  infected  by  natural  channels  with  the 
bacillus  of  epizootic  abortion. 


Schroeder  (12)  carried  out  investigations  to  ascertain  the  fre- 
quency \\'ith  wliicli  bulls  react  to  abortion  tests,  and  the  frequency 
Avith  which  lesions  chargeable  to  al)ortion  bacilli  occur  in  the  re- 
productive organs  of  reacting  bulls.  Studies  were  also  pursued 
which  he  states  conclusively  prove  that  bulls  with  infected  repro- 
ductive organs  may  expel  abortion  bacilli  with  the  seminal  fluid. 
In  the  first  two  mentioned  investigations  325  bulls  from  a  Wash- 
ington abbatoir  were  tested,  and  slaughtered  upon  reaction.  "Ap- 
proximately ten  per  cent  of  the  bulls  reacted,  and  approxi- 
mately ten  per  cent  of  the  reacting  bulls  showed  lesions  of  the 
reproductive  organs  from  which  abortion  bacilli  were  isolated.'* 
The  value  of  these  studies,  he  emphasizes,  lies  not  in  "  that  they 
give  us  a  measure  of  the  proportion  of  bulls  that  react  positively 
to  abortion  tests  or  the  proportion  of  reacting  bulls  that  are  car- 
riers of  abortion  bacilli,"  but  in  "  the  fact  that  they  show  that 
abortion  bacillus  disease  of  tlie  bull's  reproductive  organs  is  not  a 
whoU}'  unique  affection  which  practically  may  be  ignored,  but  an 
important  condition  that  must  be  taken  into  account  in  our  efforts 
to  combat  infectious  abortion,  since  it  has  been  proved  to  be  asso- 
ciated with  contamination  of  the  seminal  fluid."  In  discussing 
the  method  by  which  infected  bulls  transmit  the  organisms  to 
cattle,  he  believes  that  leakage  of  semen  from  the  penis,  or  vaginas  of 
cattle  after  service,  contaminates  the  food  which  subsequently  gains 
entrance  to  their  digestive  tracts.     As  the  result  of  a  series  of 

experiments,  he  states:  " the  results  fail  to  justify  in  the 

least  degree  the  assumption  that  cows  are  infected  with  abortion 
bacilli  via  their  vaginas  or  uteruses  at  the  time  of  copulation,  or 
that  the  bull,  through  copulation,  is  an  agent  in  the  spread  of 
abortion  disease." 

The  work  so  far  alluded  to,  has  been  limited  to  infection  with, 
and  the  transmission  of,  Bart,  ahortnm  and  the  lesions  associated 
with  such  infection.  The  last  mentioned  author,  however,  states : 
"A  search  for  other  specific  causes  of  al)ortions  among  cattle 
has  not  been  neglected,  and  bureau  investigators  could  relate  at 
great  length  stories  similar  to  those  which  investigators  have  told 
about  microorganisms  isolated  from  the  products  of  abortions  and 
the  uteruses  of  cows  that  have  aborted.  Bacilli  of  various  kinds, 
different  types  of  micrococci,  and  spirilla  or  vibrio  have  been  found 
repeatedly ;  but  when  their  pathogenicity  has  been  tested  in  accord- 
ance with  widely  recognized  and  accepted  and  required  bacterio- 
logical standards,  not  one  shred  of  evidence  has  been  obtained  to 
prove  them  true  etiological  factors  of  bovine  abortions.  What  role 
such  microorganisms  may  have  as  causes  of  the  sequellae  of  in- 
fectious abortions,  and  of  other,  possibly,  independent,  abnormal 
processes  in  the  reproductive  organs,  is  far  from  clear  and  merits 
careful  study."  Iladley  (5)  mentions  the  fact  that:  "  Unques- 
tiona1)]y  the  male  often  ])ecomes  infected  with  the  germs  that 
produce  the  various  secondary'  diseases  in  the  female,  Avhich  are 
properly  classed  under  the  more  inclusive  term  'abortion  dis- 
ease.' "    Also,  speaking  of  the  rarity  with  which  the  bull  acquires 


abortion  infection,  he  alludes  to  the  i'aet  that  he  may  act  as  a 
"  meclianical  carrier  of  various  disease  germs  from  an  infected  to 
a  healthy  cow."  Carpenter  (9)  working  on  the  female  genital  tract, 
comes  to  the  conclusion  that,  in  all  probability,  the  genital  organs 
are  normally  free  from  bacteria.  Barney  (13)  quoting  Huet  finds 
that  bacteria  may  be  present  in  the  seminal  vesicles  of  healthy 
animals  (horses,  cattle,  pigs,  and  laboratory  animals).  This,  he 
states,  corresponds  with  the  well  recognized  ilndings  in  other  parts 
of  the  genito-urinary  tract,  not  only  in  animals,  but  in  man.  He 
(Huet)  has  further  found  that  in  animals  dying  of  acute  septi- 
cemia, the  specific  organism  (anthrax,  i)neumococcus)  is  to  be 
found  in  the  vesicular  secretions.  Furthermore  it  was  definitely 
shown  that  an  infection  could  be  transmitted  to  the  female  during 
the  act  of  copulation. 

Williams,  AV.  L.  (14),  calls  attention  to  the  lack  of  vet- 
erinary literature  relating  to  the  pathology  and  bacteriology 
of  the  male  genital  tract,  except  as  related  to  infection  with 
Bad.  ahortum.  Infection  with  other  types  of  bacteria  is  empha- 
sized,  the  clinical  recognition  of  such,  with  the  accompanying 
pathological  changes,  and  of  the  numerous  phenomena  involved  in 
tlie  process  of  reproduction  in  the  male.  The  semen  and  its  essen- 
tial germinal  elements  are  taken  up  with  reference  to  the  entire 
lack  of  study  devoted  to  them,  and  some  of  the  abnormal  changes 
are  described.  In  a  later  contribution  (15),  he  takes  up  the  part 
played  by  the  bull  in  the  dissemination  of  genital  infections  and 
.states:  "  Clinical  studies  now  indicate  Avitli  great  clearness  that 
the  bull  is  an  active  spreader  of  that  group  of  genital  infections 
which  cause  sterility,  abortion,  and  related  phenomena." 

Williams,  W.  W.  (16)  studied  the  semen  with  reference  to  steril- 
ity, emphasizing  the  importance  af  its  examination  in  the  diag- 
nosis, giving  methods  for  collecting  samples,  staining'  of  sperms, 
and  some  of  the  abnormalities  encountered.  The  work  is  funda- 
mental, and  shouhl  be  of  great  practical  impoi-taiice  to  all  interested 
in  the  proljlem.  In  a  later  paper  (17),  he  brings  out  a  more  ex- 
tended discussion  of  the  question.  He  concludes  that  the  clinical 
examination  is  of  vital  importance.  v.r\(\  that  the  efficiency  of  the 
semen  depends  not  only  upon  its  physical  properties  Init  upon  thc- 
number  of  spermatozoa  that  are  motile,  the  degree  of  motility, 
degree  of  obligospermia,  and  the  percentage  of  imperfect  spermato- 
zoa, either  deformed  or  immature.  Of  forty  bulls  examined,  he 
finds  that  twenty,  or  fifty  per  cent,  sliowed  lessened  fertility,  and 
others,  aside  from  this,  showed  minor  clianges  in  the  genital 
organs  or  semen.  The  same  author  suliseciuently  takes  up  the  sub- 
ject of  reproduction  from  the  viewpoint  of  l)oth  sexes,  but  emplui- 
sizing  infection  in  the  male,  and  the  frequency  with  which  lowered 
A'itality  of  the  germinal  cells  occurs.  Hopper  (18)  states:  "A 
diseased  bull  may  manifest  non-fertility  or  decreased  potency  in 
diiTerent  ways  —  by  repeated  service  to  apparently  normal  females 
without  conception,  by  a  high  a])ortion  rate  in  females  that  have 
been  apparently  normal,  by  characteristic  infections  following!  the 


use  of  any  particular  sire,  or  by  abnormalities  in  the  breeding 
tract  noted  by  rectal  or  physical  palpation." 

The  observations  of  Williams  (19)  in  a  pure  bred  dairy  herd 
bring  out  quite  clearly  the  relation  of  the  bull  to  the  dissemination 
of  genital  infections.  The  bulls  in  this  particular  herd  were  ab- 
normal in  many  respects,  as  demonstrated  by  pathological  changes 
in  their  genital  organs,  bacterial  invasion  of  the  parts,  abnormali- 
ties of  the  semen  and  spermatozoa,  and  the  probable  transmission 
of  infection  to  the  females.  Several  of  the  sires  from  this  herd 
furnished  much  of  the  material  for  the  early  basic  work  of  this 
investigation.  Since  then  the  tracts  of  other  sires  have  been  worked 
upon  with  quite  similar  or  identical  results. 

To  summarize  the  work  already  done,  most  investigators  have 
considered  the  bull  as  merely  a  mechanical  carrier  of  Bact.  abor- 
tum  infection,  though  all  are  more  or  less  suspicious  of  his  ability 
to  become  an  active  spreader.  Schroeder,  however,  states  that  the 
organisms  are  eliminated  Avith  the  semen,  but  infection  of  the  fe- 
male occurs  secondarily  through  the  digestive  tract  by  contamina- 
tion of  the  food  with  the  semen.  Other  investigators  bring  out 
fundamental  points  demonstrating  the  importance  of  other  organ- 
isms than  the  Bang  bacillus  and  call  attention  to  the  need  of  a 
more  thorough  study  of  the  anatomy,  physiology,  and  pathology 
of  t.he  male  genital  tract. 

Any  study  of  the  genital  organs  must  of  necessity  rest  funda- 
mentally upon  a  thorough  knowledge  of  the  anatomy  and  phy- 
siology of  those  parts.  Too  few  of  us  have  stopped  to  consider 
thei^e  questions  thoroughly,  with  the  result  that  our  ideas  on  the 
problem  are  more  or  less  vague.  It  is  much  easier  to  understand 
why  abnormal  spermatozoa  occur  so  frequently,  or  changes  take 
place  in  the  semen  with  death  or  weakening  of  the  germinal  ele- 
ments, if  we  realize  or  stop  to  consider  the  highly  differential  pro- 
cess of  spermatogenesis,  and  the  various  structures  which  contrib- 
ute to  the  formation  of  the  semen.  AVe  must  come  to  realize  that 
each  part  of  the  genital  tract  is  essential  to  the  normal  functioning 
of  the  whole,  and  that  the  genital  tract  and  rer)roduction  are  in 
turn  dependent  upon  the  proper  functioning  of  the  entire  body. 

Walker  (20)  emphasizes  the  importance  of  a  thorough  knowledge 
of  physiology'  in  stating,  ''Although  the  subject  of  sterility  has 
attracted  the  attention  of  the  medical  profession;  and  although 
much  has  been  written  on  its  causes  and  treatment,  it  cannot  be 
claimed  that  the  practical  results  olitaincd  up  to  the  present  time 
are  satisfactory,  or  that  when  consulted  for  sterility,  the  medical 
man  of  today  can  hold  out  to  his  patient  much  more  hope  of  suc- 
cessful treatment  than  the  medical  man  of  fifty  years  ago.  Our 
failure  in  this  respect  is  in  the  main  due  to  an  ignorance  of  the 
physiology  of  reproduction." 

To  bring  out  some  of  these  points,  the  anatomy  of  the  tract  will 
l)e  reviewed  briefly,  together  with  the  physiology  of  reproduction, 
and  the  various  factors  which  should  be  considered  in  a  studv  of 
the  problem. 


Anatomy  and  Physiology 

In  origin  and  early  development  the  ovary  and  testis  are  identi- 
cal. The  gonad  and  mesonephros  or  primitive  kidney  are  de- 
veloped from  the  urogenital  fold.  The  gonad  first  forms  as  a 
medio-ventral  thickening  of  the  fold,  which  gradually  expands 
until  it  becomes  attached  by  a  mere  stalk.  At  first,  the  gland  is 
made  up  merely  of  a  superficial  epithelial  layer,  and  an  inner 
epithelial  mass,  or  epithelial  nucleus.  In  the  process  of  develop- 
ment, large  primordial  germ  cells  migrate  from  the  entoderm  of 
the  future  intestinal  canal,  and  pass  through  the  stalk  to  the  gonad. 
In  the  case  of  the  male  gonad,  seminiferous  tubules  are  very 
difficult  to  make  out  in  eml)ryos  smaller  than  24  millimeters.  Then 
they  suddenly  differentiate  out  as  solid  cords  of  germ  cells,  -while 
the  connective  tissue  grows  in  around  them.  These  connective 
tissue  sheaths  unite  at  the  center  of  the  organ  to  form  the  anlage 
of  the  mediastinum  testis.  The  testicular  tubules  unite  and  con- 
verge toward  the  hilus,  there  to  meet  the  anlage  of  the  rete.  At 
the  mesonephric  end  of  the  testis,  the  rete  first  appears  as  a  col- 
lection of  cells,  differentiating  out  from  the  inner  epithelial  mass 
of  the  gonad.  These  cells  gradually  grow  out  to  meet  the  collect- 
ing portions  of  the  mesonephric  tubules  on  the  one  hand,  and  the 
seminiferous  tubules  on  the  other.  The  rete  is  represented  as 
cords  of  cells  at  first,  which  in  forty  millimeter  embryos  hollow  out 
to  form  tubules. 

The  mesonephros,  or  primitive  kidney,  ea;'ly  starts  to  degenerate 
cranio-caudally, —  the  tubules  becoming  separated  into  a  cranial 
and  caudal  group.  The  collecting  and  secretory  parts  of  the 
cranial  group  separate,  the  collecting  tubules  growing  out  to  meet 
the  rete  with  wliieh  they  unite  to  form  the  efferent  ductules  of  the 
epididymis.  The  caudal  group  of  tubules  is  vestigial  and  becomes 
the  paradidymis.  The  mesonephric  duct  becomes  the  vas  deferens, 
connecting  as  it  does  with  the  tubules  of  the  epididymis,  and 
emptying  into  the  urethra  at  Mliller's  tubercle  or,  as  it  later  be- 
comes, the  colliculus  seminalis. 

The  seminal  vesicles  arise  as  hollow  saccules  from  the  dorsal 
wall  of  the  mesonephric  duct  just  as  it  empties  into  the  urethra. 
The  prostate  develops  aa  an  outgrowth  of  the  dorsal  urethra  .just 
posterior  to  ]\[iiller's  tubercle.  The  bulbo-urethral  glands  appear 
as  solid,  paired,  epithelial  outgrowths  from  the  entoderm  of  the 
urogenital  sinus. 

!Miiller's  duct,  at  first  a  solid  tube  growing  from  the  anterior 
part  of  the  mesonepliros,  and  ending  at  ^Midler's  tubercle,  becomes 
a  hollow  tube,  and  in  the  female  forms  the  entire  genital  tract  ex- 
cept for  the  gonad  and  the  lower  part  of  the  vagina.  In  the  male, 
the  anterior  part  remains  as  the  vestigial  appendix  testis,  and  the 
posterior  part,  as  the  vagina  masculina,  Ellenberger  states,  how- 
ever, that  this  embryonic  structure  is  very  seldom  seen  in  the 
mature  bull. 


10 

The  Male  Reproductive  Organs  include  the  penis  and  testes, 
together  with  the  excretory  passages  which  connect  the  testes  with 
the  urethral  canal.  These  excretory  duets  include  the  epididymis, 
vas  deferens,  and  seminal  vesicles.  Posterior  to  their  termination 
in  the  urethra,  there  aro  connected  the  ducts  of  the  prostate  gland 
and  the  bulbo-urethral  or  Cowper's  glands. 

Testes  :  The  testicles  of  the  bull  are  relatively  large.  Vaiying 
with  the  size  and  age  of  the  animal  they  measure  from  fourteen  to 
seventeen  centimeters  in  length,  including  the  epididymis,  and 
from  six  to  eight  centimeters  in  diameter.  Each  testicle  is  en- 
closed within  a  serous  sac,  the  tunica  vaginalis,  whose 
visceral  layer  is  very  intimately  fused  with  the  underly- 
ing covering  of  the  organ,  the  tunica  albuginea.  The  tunica 
albuginea  is  quite  thin  and  consists  of  connective  tissue 
which  is  rich  in  elastic  fibres.  Muscular  tissue  is  not  present  as  it 
is  in  the  case  of  many  mammals.  Inside  the  tunica,  and  closely 
attached  to,  though  separated  from,  the  parenchyma  by  a  thin 
layer  of  connective  tissue,  is  a  laj-er  of  very  loose  connective 
tissue,  M'hich  because  of  its  rich  supply  of  blood  vessels  is  termed 
the  tunica  vasculosa.  The  parenchyma  is  of  a  yellowish  gray 
color,  and  of  a  rather  soft  consistency.  It  is  made  up  of  the  semi- 
niferous tubules,  rete,  and  the  connective  tissue  stroma,  the  medi- 
astinum testis.  On  section,  the  mediastinum  appears  as  the  center 
or  axis  of  the  entire  organ.  It  is  star-shaped,  and  radiates  connect- 
ive tissue  septa  out  into  the  parenchyma  to  support  and  separate  the 
tubule.^.  Ellenberger  states  that  the  testis  of  the  bull  and  all 
ruminants  lacks  a  closed  system  of  interlobular  septa,  because  of 
the  feeble  development  of  the  connective  tissue. 

The  principal  blood  vessels  and  rete  tubules  are  found  in  this 
structure,  the  function  of  the  latter  being  to  connect  the  seminifer- 
ous tubules  and  the  efferent  tubules  of  the  epididymis.  The 
epithelium  of  the  rete  is  quite  irregular. —  consisting  in  places  of 
a  single  layer ;  in  others,  of  two  layers.  At  some  points  there  are 
formed  groups  of  several  cells  lying  over  one  another,  with  swollen 
liomogeneous  basal  cells,  which  sometimes  form  projections  into  the 
lumen. 

The  interstitial  tissue,  besides  conveying  the  blood  vessels  to  the 
organ,  contains  many  "  interstitial  cells.'-'  These  cells  are  rela- 
tively sparse  in  the  adult  bull,  and  are  comparatively  delicate, 
slightly  granular,  often  shuttle-shaped,  with  a  rather  small  nucleus. 
Embryologicnlly  they  are  derived  from  a  syncitium  arising  from 
the  mesothelium  of  the  genital  ridge,  differentiating  out  by  growth 
of  the  cytoplasm.  They  contain  large  quantities  of  fat,  and 
elaborate  the  internal  secretion  of  the  testis.  This  secretion  gov- 
erns the  development  of  the  secondary  sexual  characters,  and  has 
a  profound  influence  on  the  general  body  metabolism,  and  develop- 
ment of  the  skeleton.  The  interstitial  cells  appear  early  in 
embryonic  life  even  before  there  is  any  differentiation  of  sex,  and 


11 

their  greater  relative  developmeut  iu  the  fetus  is  indicative  of  a 
future  male  development.  In  very  young  embryos,  the  growth  is 
very  rapid,  followed,  however,  by  a  period  of  atrophy,  during 
which  the  seminiferous  tubules  undergo  marked  development. 
Pende  (21)  states:  "  There  seems  to  be  an  inverse  relation  be- 
tween the  growth  of  the  tubular  and  interstitial  tissues,  as  one  is 
hypoplastic  when  the  other  is  in  full  activity."  From  birth  to  the 
onset  of  sexual  maturity,  whicli  may  be  called  a  period  of  rest  for 
the  testicle,  the  cells  are  few  in  number.  With  the  accentuation  of 
the  secondary  sexual  characters,  and  the  beginning  of  sexual  life, 
these  cells  again  increase  in  number  and  acti^-ity. 

The  parenchyma  of  the  testis  consists  for  the  most  part  of  the 
seminiferous  tubules,  which,  on  account  of  the  courses  they  take 
in  the  different  regions,  are  divided  into  groups.  The  peripheral 
tubules  are  the  much-contorted  tubuli  contorti.  These  anastomose 
to  form  the  mucli  shorter  tul)idi  recti.  These  in  turn  anastomose 
frequently,  uniting  to  form  the  rete  testis.  The  rete  proceeds 
through  the  mediastinum  to  form  the  efferent  ductules  which 
break  through  the  tunica  albuginea  to  form  the  greater  part  of  the 
liead  of  the  epididymis.  The  tubuli  contorti  are  the  longer  and 
more  numerous  of  the  tubules,  for  it  is  liere  that  practically  all 
the  spermatozoa  are  produced.  The  straight  tubules  are  relatively 
so  short  that  they  may  be  regarded  more  in  the  light  of  the  begin- 
ning of  the  system  of  excretory  ducts. 

The  seminiferous  tubules  consist  of  a  thin  peripheral  meml)rana 
l)ropria  upon  which  rests  tlie  seminal  e])ithelium,  which  is  made  up 
of  the  essential  semen  forming  cells,  and  the  cells  of  Sertoli.  The 
spermatogenic  cells  may  be  divided  into  three  groups,  from  within 
outward:  the  peri])heral  single  layer  of  small  cuboidal  spermato- 
gonia ;  one  or  two  rows  of  large  spermatocytes ;  and  three  to  five  roAvs 
of  spheroidal  spermatids.  Tlie  cells  of  Sertoli  are  more  or  less  of  the 
syncitial  type. —  large  in  size  and  irregular  in  outline.  They  occur 
at  various  intervals  between  the  layers  of  spermatogenic  cells,  with 
their  bases  resting  upon  the  mcmbrana  proi)ria.  Centrally  they 
send  out  protoplasmic  processes  for  variable  distances,— some 
even  reaching  the  l)order  of  the  innermost  cell  layers. 

Spermatogenesis:  In  this  process,  the  ])rimary  germinal  cells, 
the  spermatogonia,  divide  to  form  the  i)riniary  spermatocytes, 
^laturation  consists  of  two  cell  divisions  of  tlie  primary  sperma- 
tocytes, and  these  in  turn  form  four  spermatids.  During  the  pro- 
cess, the  number  of  chromosomes  is  reduced  to  half  the  number 
characteristic  of  the  species.  The  spermatids  then  become  con- 
verted into  mature  spermatozoa.  Tliis  mode  of  transformation 
may  be  seen  in  Plate  I.  In  the  process,  the  nucleus  of  the  sperma- 
tid forms  a  large  part  of  the  head;  the  eentrosome  divides,  part 
passing  to  the  extremities  of  the  neck.  One  eentrosome  ])ecoraes 
the  .interior,  and  remains  altaelied  to  the  head,  while  tlie  other 
divides  to  form  the  posterior  eentrosome.  The  latter  is  divided 
into  the  anterior  part,  and  the  posterior  nodule  or  annular  ring. 


12 

Besides  this,  the  posterior  eentrosome  becomes  elongated  to  form 
the  axial  filament,  and  the  cytoplasm  forms  the  sheaths  of  the  neck 
and  tail.  The  spiral  fihiment  of  the  connecting  piece  is  derived 
from  the  cytoplasmic  mitochondria.  At  this  time,  a  large  part  of 
the  cellular  cytoplasm  is  cast  off.  Meanwhile,  the  spermatozoa 
sink  their  heads  into  the  long  protoplasmic  processes  of  the  Sertoli 
or  "  nurse  "  cells  which  furnish  nutritive  material  for  their  com- 
plete development.  Finally  the  adult  cells  are  cast  off  into  the 
lumen.  The  structure  of  the  spermatozoa,  and  a  discussion  of  the 
semen  will  be  taken  up  later. 

Epididymis:  The  epididymis  is  divided  into  three  parts:  the 
head,  body  and  tail.  The  head  is  made  up  principally  of  the 
lobules  formed  by  the  much-coiled  efferent  ductules  proceeding 
from  the  rete.  The  ductules,  about  twelve  in  number  in  the  bull, 
unite  to  form  the  body,  which  remains  coiled  and  runs  along  the 
postero-medial  part  of  the  testicle  to  which  it  is  more  or  less  closely 
attached.  To  quote  Ellenberger  (23)  :  "  The  transition  from  the 
rete  into  the  ductules  is  gradual,  as  the  characteristic  epithelium 
of  the  latter  (ductules)  begins  in  cavities  without  walls,  and  at 
first,  gradually  form  a  wall  which  is  well  marked  out  as  a  thin 

ring  of  interstitial  tissue The  epithelium  of  the  ductules  is 

in  sharp  contrast  to  the  rete  in  that  it  has  a  single-layered  ciliated 
columnar  epithelium,  in  which  here  and  there  one  finds  round  basal 
cells.  The  dark  and  light  columnar  cells  alternate;  the  cilia  are 
often  cemented  together,  and  form  cone-shaped,  homogeneous 
appearing  protuberances.  The  secretorj^  activit}^  is  quite  clearly 
observable.  In  the  light  cells  one  finds  secretory  globules,  accumu- 
lating in  rows,  sometimes  above,  other  times  below,  arches  of  cells. 
The  secretory  droplets  pass  from  the  cells  into  the  lumen,  and 
often  lie  in  irregular  layers  on  the  epithelium ;  also  the  basal  cells 
appear  swollen  and  shoved  out  between  the  cylindrical  cells."  At 
the  lower  extremity  of  the  testicle,  the  tail  is  formed,  which  is 
globular  in  shape,  and  more  or  less  loosely  attached  to  the  testicle. 
Here  the  ductules  anastomose  freely,  gradually  become  less  coiled, 
and  end  in  the  single  excretory  tube,  the  vas  deferens.  The  epithe- 
lium at  the  tail  part  is  more  or  less  of  the  pseudo-stratified 
columnar,  ciliated  variety.  Outside  this  is  a  membrana  propria, 
a  circular  muscular  layer,  and  a  connective  tissue  coat.  The  secre- 
tory activity  is  very  marked  here  and  one  finds  much  secretion  in 
the  lumen.  Courrier  (24),  working  on  the  bat,  suggests  that  the 
glandular  activity  is  conditioned  by  the  secretion  from  the  inter- 
stitial (endocrine)  gland.  The  action  of  the  secretion  is  to  dilute 
the  large  mass  of  spermatozoa  present,  nourish  them  to  some  extent 
and  also  stimulate  them  to  active  motility.  Stigler  (25)  states 
that  the  properties  of  the  sperms  are  modified  in  the  epididymis; 
the  motility,  the  ability  to  resist  heat,  and  other  properties  are 
augmented,  at  least  in  the  case  of  the  guinea  pig,  rat,  and  mouse. 
Some  authors  state  that  the  sperms  first  become  motile  when  in 
contact  with  the  prostatic  secretion,  but  I  have  repeatedly  exam- 


13 

ined  the  contents  of  the  tail  of  the  epididymis  of  the  bull,  rabbit, 
and  guinea  pig,  finding  full  motility  in  each  case,  though  the  dura- 
tion is  not  nearly  as  long  as  when  the  sperms  are  ejaculated  in  the 
semen. 

The  Vas  Deferens  is  quite  narrow  (2  mm.)  and  runs  from  the 
tail  of  the  epididymis  to  the  verumontanum,  or  colliculus  semin- 
alis,  w'here  it  empties  into  the  urethra  in  common  with  the  duct  of 
the  vesicle.  At  first  it  is  lined  by  epithelium  similar  to  that  of  the 
vas  epididymis,  but  this  changes  over  into  a  peculiar  low  stratified 
type.  Ellenberger  describes  it  as  follows:  "  The  epithelium 
shows  a  very  pronounced  basal  coat.  The  overlying  cell  zone 
shows  more  (at  the  most,  three)  rows  lying  over  each  other  of 
elongated  nuclei,  while  an  outline  of  cell  form  is  not  ordinarily 
noticeable,  so  that  it  may  be  spoken  of  as  a  syncitium,  and  at  the 
same  time  as  a  many  layered  epithelium."  The  mucosa  forms  low, 
broad  folds  into  the  lumen.  The  tunica  propria  is  a  thin  con- 
nective tissue  layer.  The  submucosa  consists  of  thin  connective 
tissue.  Three  muscular  coats  are  present :  an  inner  thin  longitud- 
inal layer,  middle  circular  layer,  and  an  outer  longitudinal  layer. 
All  are  more  or  less  intimately  blended,  and  are  surrounded  by 
the  adventitia,  made  up  of  connective  tissue,  elastic  fibres  and 
scattered  longitudinal  muscle  cells  of  the  internal  cremaster  mus- 
cle. Near  the  dorsal  siu*face  of  the  bladder,  the  ducts  come  in 
close  apposition,  and  for  ten  to  twelve  centimeters  dilate  to  form 
the  ampullae.  Here  the  mucous  membrane  bcomes  much  plicated, 
forming  long  folds  which  anastomose  freely.  The  function  of  the 
vas  is  to  convey  the  spermatozoa  and  secretions  from  the  epididymis 
to  the  urethra.  Disselhorst  (26)  believes  the  ampulla  acts  "as  a 
seminal  reservoir  and  states  that  he  has  found  spermatozoa  stored 
up  in  the  little  pockets  in  the  walls  of  this  structure  in  animals 
during  the  rutting  time.  He  suggests,  further,  that  there  is  a 
relation  between  the  state  of  development  of  the  ampulla  and  the 
time  occupied  by  copulation.  When  the  organ  is  small  or  absent, 
as  in  dogs,  cats,  and  boars,  the  coition  is  a  slow  process,  but  when 
the  ampulla  is  large  and  well  developed,  as  in  horses  and  sheep, 
the  coitus  requires  a  relatively  short  time.  Inasmuch  as  coitus  is 
so  rapid  in  the  bull,  and  the  ampulla  is  so  well  developed,  it  seems 
as  though  this  function  is  very  probable. 

The  Seminal  Vesicles  are  very  compact  glandular  structures 
^ylng  on  either  side  of  the  median  line,  on  the  dorsal  side  of  the 
'ladder,  and  ventral  to  the  rectum.  In  the  mature  bull  they 
rlieasure  ten  to  twelve  centimeters  in  length,  four  centimeters  in 
width,  and  about  two  and  one-half  to  three  centimeters  in  thick- 
ness. The  glands  are  distinctly  lobulated.  quite  tortuous,  and  are 
often  asymmetrical  in  size  and  shape.  They  converge  posteriorly, 
to  empty  into  the  urethra  at  the  colliculus  seminalis  with  the 
ampulla,  in  a  slightly  oval  slit  in  the  mucosa.  IMicroscopically, 
the  gland  is  of  the  anastomosing  tubular  type,  with  very  poorly 


14 

developed  excretory  ducts  to  the  glandular  cavities.  Posteriorly 
one  finds  centrally  a  few  sinus-like  narrow  excretory  passages, 
which  open  into  the  somewhat  larger  collecting  and  excretory  duct. 
The  epithelium  is  of  the  simple  columnar  tj-pe  and  produces  a 
relatively  large  amount  ol'  secretion.  The  gland  cavities  are  sur- 
rounded by  a  membrana  propria,  over  which  is  a  relatively  thick 
layer  of  smooth  muscle.  Outside  this  is  a  connective  tissue  covering 
which  sends  traljeculae  or  septa  in  between  the  lobules.  The  secre- 
tion of  the  seminal  vesicles  is  a  tenacious  albuminous  tliiid  with 
a  slightly  yellowish  tinge,  all  or  part  of  which  appears  in  the 
ejaculate  in  the  form  of  swollen  sago-like  grains  which  are  soon 
dissolved  following  ejaculation  and  the  liquefaction  of  the  semen. 
The  proteid  compounds  belong  to  the  group  of  histones.  The 
secretion  is  liquid  when  warm  and  coagulates  when  cold.  Some 
say  that  the  filling  of  the  vesicles  serves  to  excite  sexual  feeling, 
but  this  is  doul)tful  in  view  of  the  fact  that  in  some  animals  the 
sexual  desire  exists  before  the  vesicles  are  filled.  Likewise,  Stein- 
ach  found  that  rats,  whose  seminal  vesicles  had  been  removed, 
still  retained  their  desire  for  copulation.  The  function  of  the 
secretion  is  to  furnish  much  of  the  volume  to  the  semen,  and  in 
some  way  it  has  a  distinct  1)earing  on  fei1:ility,  inasmuch  as  ex- 
tirpation of  the  organs  in  rats  leads  to  lowered  fertility.  The 
vesicles  of  the  bull  are  in  no  sense  a  store-house  for  spermatozoa, 
as  is  usually  understood.  Repeated  examinations  in  a  large  num- 
ber of  bulls  have  led  to  the  finding  of  spermatozoa  there  only  in  veiy 
rare  instances.  That  they  serve  as  a  resorption  j^lace  for  sperms 
that  are  not  ejaculated  is  also  very  unlikely.  Normally,  one  sees 
on  smear  of  the  vesicles,  occasional  cells,  leucocytes,  lecithin  gran- 
ules, sago  bodies,  and  rarely  a  few  degenerated  spermatozoa. 

The  CoLLicuLUS  Se:mixalis  is  a  rounded  or  cone-shaped  emi- 
nence in  the  posterior  urethra,  upon  which  the  ducts  of  the  sem- 
inal vesicles  and  vasa  deferentia  open.  The  ducts  open  separately 
at  the  bottom  of  two  narrow  slits,  one  on  each  side  of  the  mound, 
there  being  no  distinct  ejaculatory  duct  as  in  man.  The  function 
of  the  eollicuius  or  veruinontanum  is  not  definitely  known.  It 
is  generally  believed  that  the  structure  is  made  up  of  blood  spaces 
which  become  engorged  during  erection,  causing  a  l)lockage  of 
the  posterior  urethi'a.  ^»•hich  jirevents  regurgitation  of  the  semen. 
Iivtina  (27),  howcNcr,  demonstrated  tliat  the  structure  is  not  com- 
posed of  any  unusual  number  of  blood  vessels  or  spaces,  and  that 
removal  of  the  organ  Avas  not  followed  by  regurgitation  of  the 
semen  into  the  l)ladder  during  ejaculation.  Tie  ])elieves,  and  quite 
logically,  that  its  function  is  to  afford  a  prominence  upon  which 
the  duets  may  empty.  The  mixture  of  the  thick  gelatinous  semen 
with  the  tliin  ])rostatic  secretion  must  occur  at  the  moment  of 
ejaculation  and  must  be  jierfectly  homogeneous,  otherwise  large 
num])ers  of  the  organisms  remain  in  the  thick  gelatinous  portion 
of  the  fluid.  The  eminence  serves  this  purpose  in  that  the  pros- 
tatic ducts  which   converge  toward   it,   may  eject  their  secretion 


15 

toward  the  eminence,  producing  an  admixture  more  evenly  and 
quickly. 

Prostate:  The  hull  possesses  -what  Ellenherger  calls  a  diffuse 
prostate.  That  is,  there  is  no  distinct  glandular  hody  as  in  man. 
It  is  composed  principally  of  a  glandular  sheath  around  the 
urethral  wall.  Just  posterior  to  the  neck  of  the  bladder,  and  in 
front  of  the  urethral  muscle,  there  is  formed  a  slight  dorsal  trans- 
verse elevation,  extending  downward  on  the  sides.  This  is  what 
might  be  termed  the  body.  The  greater  part  of  the  gland  is 
"disseminate"  in  form,  being  a  sheath  of  glandular  tissue  em- 
bedded in  the  urethral  wall.  Dorsally  it  is  about  ten  to  twelve 
millimeters  thick,  and  ventrally  about  two  millimeters.  The  gland 
is  a  branched  tul)ular  structure,  the  interlobular  tissue  of  which 
contains  much  unstriped  muscle.  The  lobules  arc  lined  by  a 
columnar  type  of  epithelium.  The  ducts,  about  thirty  to  forty 
in  numl)er,  o])en  into  the  urethra  in  two  rows  posterior  to  the 
colliculus.  The  secretion  is  a  thin,  slightly  turbid  fluid,  of  a 
faintly  alkaline  reaction.  Its  function  is  to  dilute  the  semen, 
stimulate  the  motility  of  the  spermatozoa  and  nourish  them. 

Fish  (28)  believes  that  the  activating  property  of  the  secretion 
is  due  to  enzymes,  because  boiling  deprives  the  fluid  of  its  power 
to  accelerate  the  motility  of  the  spermatozoa.  Serrlach  and  Pares, 
quoted  by  ]\rarshall  (29),  working  on  dogs,  have  adduced  evidence 
indicating  that  the  prostate  is  an  internal  secretory  gland  which 
controls  the  testicular  functions,  and  regulates  the  process  of 
ejaculation.  It  is  stated  that  if  the  prostate  is  removed,  sperma- 
tozoa are  no  longer  produced  in  the  testis,  and  that  the  secretory 
activity  of  the  accessory  genital  glands  ceases.  These  changes, 
however, 'can  be  prevented  by  the  administration  of  extracts  of 
the  prostate.  The  fact  that  the  prostate  elaborates  a  secretion 
having  a  definite  relation  to  the  testis,  has  been  verified  by  other 
authors. 

Cowper's  Geaxds  (Bull)o-Urethral)  :  These  glands  are  paired, 
oval  structures  about  one  and  one-half  by  two  and  one-half  centi- 
meters in  size,  situated  on  either  side  of  the  dorsal  pelvic  part  of 
the  urethra  close  to  the  ischial  arch.  They  are  deeply  embedded, 
with  the  bull)us  urethae,  in  the  bulbo-cavernosus  muscle,  thereby 
being  inaccessil)le  to  palpation.  Tn  general,  they  are  of  a  well 
developed  anastomosing  tubular  type.  The  connective  tissue 
.stroma  is  relatively  thin,  and  thickens  only  in  between  the  larger 
lobules,  where  one  finds  the  larger  collecting  ducts.  Each  gland 
opens  by  a  single  duct.  The  epithelium  is  of  the  sim])le  cuboidal 
type,  ijittle  is  known  of  the  function  of  its  .secretion,  though 
Kingsbury  '':"50^  regards  it  in  the  light  of  a  mating  gland;  that  is. 
it  lubricates  the  genital  passages  dui-ing  coitus,  as  does  its 
homologous  structure  in  the  female,  Bartholin's  gland. 

Ellenberger  describes  the  urethra,  .slit  ventrally,  as  presenting 
the  following  picture:     "The   colliculus  seminalis  distinctly   ap- 


16 

pears  as  a  process  or  offshoot  of  the  crista  urethralis  of  the  Tri- 
gonium  Vesicae.  At  the  summit,  and  at  the  bottom  of  the  two 
slits,  open  laterally  the  ducts  of  the  vesicles,  and  medially  the 
ductus  deferens  ....  From  the  caudal  slope  of  the  colliculus 
go  two  distinct  mucous  membrane  folds  which  run  through  the 
entire  pelvic  urethra,  near  together,  somewhat  diverging,  and  then 
coming  together,  so  that  they  form  an  elongated,  narrow  oval.  At 
their  caudal  union,  the  excretory  ducts  of  the  bulbo-urethral 
glands  open  side  by  side.  At  the  point  of  departure  of  the  folds 
from  the  colliculus,  arises  a  niche-shaped  opening,  between  both 
folds,  and  likewise  lateral  to  each  fold.  In  these  niches  open  the 
ducts  of  the  prostate.  The  openings  of  the  disseminate  prostate 
lie  in  rows  as  in  the  horse,  but  form  not  less  than  six  rows.  There 
is  one  row  medial  to  each  fold,  and  two  lateral.  Midler  mentions 
only  the  medial  rows.  These  rows  extend  to  the  opening  of  the 
ducts  of  Cowper's  glands.  The  stratum  glandulare  (disseminate 
prostate)   is  very  easy  to  recognize  with  the  naked  eye." 

Semen  :  The  semen  is  the  mixed  product  of  the  testicles,  their 
excretory  passages,  and  the  accessory  sexual  glands,  a  fact  which 
complicates  its  study  considerably.  The  freshly  ejaculated  fluid  is 
cloudy,  tenacious,  more  or  less  coagulable,  and  is  rich  in  albumen. 
It  is  weakly  alkaline  in  reaction,  and  contains  eighty  to  ninety 
per  cent  of  water.  Of  the  solid  constituents,  there  is  forty  per 
cent  of  ash,  of  which  three-fourths  is  calcium  phosphate.  Besides 
the  spermatozoa,  the  semen  frequently  contains  epithelial  cells, 
leucocytes,  concentric  amyloid  concretions,  and  lecithin  bodies. 
When'  cold,  the  characteristic  phosphoric  acid  salts  are  precipi- 
tated. The  fluid  content  is  the  product  of  the  tubules  of  the 
testicles,  their  excretory  ducts,  and  the  accessory  sexual  glands. 
The  characteristic  odor  "of  the  semen  is  supplied  by  a  slimy  nucleo- 
albumen  "spermin"  which  forms  the  spermatic  crystals,  and  is 
furnished  by  the  prostatic  secretion. 

During  ejaculation  the  spermatozoa  and  secretions  added  by  the 
testicle  and  epididymis  are  probably  carried  to  the  ampulla  by 
peristaltic  muscular  action,  in  the  earlier  stages  of  the  orgasm. 
At  the  height  of  the  orgasm,  the  ampulla  is  emptied  into  the  pos- 
terior urethra  in  common  with  the  secretion  of  the  contracting 
vesicles,  here  to  be  admixed  with  the  thin  prostatic  secretion.  The 
entire  mixture  is  then  propelled,  and  ejaculation  produced  by 
strong  muscular  contractions  of  the  entire  urethra.  As  was  stated 
before,  the  semen  is  the  product  of  the  testicles,  the  excretory 
ducts,  and  the  accessory  sexual  glands.  The  testes  furnish  the 
essential  germinal  elements,  the  sperms,  and  some  of  the  fluid  con- 
tent. Then  is  added  the  product  of  the  epididymis,  vas,  and 
ampulla,  which  stimulates  the  spermatozoa  to  active  motility, 
nourishes  the  organisms,  and  adds  somewhat  to  the  fluid  bulk  of 
the  secretion. 

Stigler  (25)  states:  "During  its  sojourn  in  the  epididymis,  the 
properties  of  the  spermatozoa  are  modified;  the   r.^vtility,  ability 


17 

to  resist  heat,  and  other  properties  are  augmented,  at  least  in  the 
case  of  the  guinea  pig,  rat,  and  mouse."  To  the  secretion  is  then 
added  the  product  of  the  vesicles,  Avhich  contributes  markedly  to 
its  fluid  content,  nourishes  the  sperms,  and  supplies  the  ferment 
which  induces  clotting  of  the  semen  when  ejaculated.  This  is 
very  important  because  the  clot  formed  in  the  vagina  protects 
the  delicate  spermatozoa  from  the  hostile  acid  vaginal  secretion. 
The  prostate  likewise  adds  bulk  and  nourishing  substances,  besides 
stimulating  the  spermatozoa  to  fuller  and  more  lasting  motility. 
The  addition  of  the  spermin  is  perhaps  unimportant.  The  func- 
tion of  the  secretion  of  the  Cowper's  glands,  which  is  added  at  this 
time,  is  problematical.  It  does,  however,  have  a  diluting  action  on 
the  semen.  Perhaps  its  secretion  is  poured  out  prior  to  ejacula- 
tion so  as  to  lubricate  the  canal  and  prepare  the  way  for  the  semen. 
Fish  (28)  has  demonstrated  by  means  of  darkfield  illumination, 
the  presence  of  numerous  minute  particles  or  ultraparticles  in  this 
fluid.  Their  character  and  significance  are  matters  of  conjecture, 
but  it  would  seem  as  though  they  were  not  identical  with  the 
"chylomicrons"  or  fat  particles  found  in  the  blood  by  Gage.  Per- 
haps further  researches  wall  reveal  some  intimate  connection  be- 
tween the  number  present  in  a  field,  and  the  relative  potency  of 
the  animal. 

Each  portion  of  the  tract  furnishes  some  essential  element  to 
the  mixed  product  which  is  so  remarkably  adapted  both  as  ^  a 
vehicle  for  the  ejaculation  of  the  spermatozoa,  and  as  a  fluid  in 
which  their  motility  is  initiated  and  maintained.  Any  derange- 
ment of  one  part  is  fraught  with  danger  to  the  existence  of  viable 
spermatozoa,  and  the  continuation  of  full  fertility  on  the  part 
of  the  animal.  The  physiology  of  each  contributing  gland  must 
be  borne  in  mind  at  all  "times.  AValker  (31)  investigated  the  fer- 
tility of  the  semen  of  the  dog,  taken  from  various  parts  of  tho 
tract.  His  results  were:  (1)  semen  from  the  testicle  and  head  of 
the  epididymis  showed  no  motility,  (2)  semen  from  the  tail  of 
the  epididymis  showed  some  motility  in  the  more  fluid  contents 
of  the  preparations,  (3)  semen  from  the  vas  deferens  appeared 
about  the  same,  (4)  a  mixture  of  epididymis  semen  and  pros- 
tatic secretion  showed  active  motility,  and  (5)  likewise  in  a  mix- 
ture of  epididymis  semen,  though  only  in  thase  places  where  the 
fluids  had  become  well  mixed.  'My  observations,  however,  differ 
in  one  respect  with  regard  to  the  Inill,  as  I  have  found  full  motility 
of  the  spermatozoa  from  the  epididymis,  but  it  is  not  so  lasting  as 
when  augmented  by  the  addition  of  the  prostatic  fluid.  Boettcher 
(32)  concludes:  "that  the  secretion  of  the  accessory  male  genital 
organs  possesses  a  protective  colloid,  which  (1)  hinders  the  sper- 
matozoid  action  of  the  vaginal  secretion,  at  least  until  the  sperms 
have  time  to  reach  the  interior  of  the  uterus  which  is  an  alkaline 
reaction,  (2)  that  it  makes  the  ejaculate  more  voluminous,  so  that 
by  cohabitation,  a  very  good  part  of  the  vagina  becomes  moistened, 
and   the   spermatozoa  become  distributed   over   the   greater   part 

2 


IS 

of  the  vaginal  mucosa.  This  distri1)ution  is  rendered  necessary 
because  some  of  tlie  fluid  flows  from  the  vagina  following  coitus. 
In  this  manner  the  opportuniiy  is  given  for  a  part  of  the  ejacu- 
late containing  the  spermatozoa  to  be  brought  easier  to  the  ex- 
ternal OS,  and  (3)  it  happens  that  because  of  its  content  of  sodium 
chloride  the  life  of  the  spermatozoa  is  stimulated  and  prolonged." 
A  fuller  discussion  of  the  essential  physiology  of  the  various  parts 
of  the  tract  on  the  semen  content,  and  fertility',  Avill  be  taken  up 
later.  The  changes  in  biochemical  content,  reaction,  and  the  re- 
sult of  the  addition  of  bacterial  products  will  also  be  more  fully 
discussed. 

Spermatozoa  :  The  history  of  the  discovery  of  spermatozoa 
is  very  interesting,  and  for  that  reason  a  brief  outline  will  be 
given.  The  "semen  threads"  were  first  observed  in  the  year 
1667,  bv  Ham,  a  student  of  Leeuwenhoek  at  Levden.  The  dis- 
covery  was  announced,  confirmed  by  findings  in  the  dog  and 
rabbit,  and  discussed  by  the  latter  author  under  the  title :  Ob- 
servationes  Anthonii  L.  de  natis  e  semine  genitali  animaculis 
(Upon  the  formation  of  young  from  procreative  material).  The 
sperms  were  taken  to  ])e  animals  on  account  of  their  motility,  and 
their  significance  remained  questionable  if  not  unknown.  Spal- 
lanzi,  quoted  by  ^Marshall  (29),  was  the  first  to  show  that  the 
filtered  fluid  was  impotent,  and  that  spermatozoa  in  the  semen 
were  essential  to  fertilization.  Kolliker,  in  1841.  discovered  that 
the  sperms  arise  from  the  cells  of  the  testis,  and  Barry  in  1843. 
observed  the  conjugation  of  sperm  and  ovum  in  the  rabbit.  This 
led  to  a  clear  understanding  of  the  function  of  these  important 
germinal  elements. 

The  spermatozoa  are  the  male  procreative  cells,  and  are  cliar- 
acterizecl  by  the  possession  of  a  head  containing  the  chromo.somes 
necessary  for  fertilization,  and  a  tail  capable  of  propelling  the 
organism  on  its  way  to  meet  the  ovum.  The  length  of  the  entire 
sperm,  including  the  head,  is  seventy-five  to  eighty  microns.  The 
head  is  nine  and  five-tenths  microns  long,  and  five  and  five-tenths 
microns  wide.  It  may  be  divided  into  two  principal  parts,  the 
head  and  tail.  The  head,  for  the  larger  part,  is  made  up  of  the 
nucleus,  and  may  be  differentiated  by  staining  reactions  into  a 
darker  staining  posterior  part,  an  anterior  lighter  part,  and  often 
a  still  lighter  area  between  the  two.  On  the  anterior  part  is  a 
sharpened  edge,  the  acrosome,  which  serves  to  perforate  the  ovum. 
The  Avhole  is  surrounded  by  a  very  definite  limiting  membrane 
which  often  becomes  obscured  under  abnormal  conditions.  The 
tail  may  be  divided  into  three  parts:  connecting  piece,  principal 
part,  and  terminal  filament.  The  connecting  piece,  the  essential 
motile  apparatus,  is  the  thickest  and  strongest  part,  and  joins  the 
tail  p-roper  to  the  head.  It  consists  of  the  central  axial  filament,  a 
spiral  filament  around  this,  and  an  outer  mitochondrial  coverinsr. 
Anteriorly  it  is  limited  by  the  anterior  portion  of  the  posterior 
chromosome,   and   posteriorly  liy  the   annular   chromosome.     The 


19 

anterior  chromosome  is  directly  connected  with  the  head,  there 
appearing  a  light  break  here  at  the  neck  where  separation  fre- 
quently occui-s.  This  neck  ser^-es  as  a  joint  for  the  motion.  The 
axial  filament,  therefore,  does  not  reach  the  head,  but  extends 
back  from  the  anterior  part  of  the  posterior  chromosome.  The 
principal  part  consists  merely  of  the  axial  filament,  and  a  thin 
outer  covering,  while  the  end  piece  is  quite  thin  and  is  made  up 
solely  ol  the  uncovered  axial  filament.  The  finer  structures  are 
seen  only  when  special  staining  reactions  are  used,  and  then  only 
when  the  sperms  are  obtained  directly  from  the  testicle.  The 
function  of  the  sperms  is  of  course  primarily  that  of  fertilization. 
Numerous  observers  have,  however,  thought  that  they  might  have 
some  other  definite,  thougli  unknown,  use. 

An  editorial  in  the  Journal  of  the  American  Medical  Associa- 
tion (33)  raises  several  important  questions  regarding  this  ob- 
scure phenomenon.  The  fact  that  an  enormous  number  of  spenna- 
tozoa  are  produced,  and  only  one,  or  at  mo.st,  a  few  perform  the 
function  of  fertilization,  raises  the  question  as  to  what  becomes 
of  the  remainder.  It  is  stated:  "Zoologists  have  found  that  in 
some  of  the  invertebrates  the  spermatozoa  invade  the  entire  body 
of  the  female,  and  in  some  species  they  reach  the  ovum  by  pene- 
trating the  cuticle  from  outside  and  migrating  to  their  goal. 
Studies  on  rodents  show  that  the  sperms  invade  the  epithelium  of 
the  generative  mucosa  and  underlying  connective  tissue.  These 
tissues  seemed  to  be  stimulated  to  growth,  suggesting  that  this 
may  influence  the  uterine  mucosa  in  its  preparation  for  receiving 
and  embedding  the  egg,  and  in  forming  the  decidua."  It  has 
been  shown  that  the  sperms  contain  a  specific  protein  capable  oC 
producing  antibodies  in  the  blood  plasma,  by  citing  the  fact  that 
rabbits  develop  a  distinct  Aberhalden  reaction  for  testicular  pro- 
teins shortly  after  cohabitation.  One  very  important  observation 
showed  tliat  by  immunizing  female  rabbits  with  sperms  they  were 
rendered  sterile  for  some  time,  although  after  a  few  months  they 
again  became  capable  of  impregnation.  The  question  raised  is: 
**....  if  the  spermatozoa  invade  the  female  tissues  and  cause 
the  formation  of  specific  antibodies  Avhich  are  capable  of  prevent- 
ing fertilization,  may  not  such  a  process  participate  in  the  prob- 
lem of  sterility?"  This  very  problem  seems  to  be  a  factor  in  ex- 
plaining why  some  couples  who  are  not  fertile  to  each  other 
subsequently  are  both  fertile  when  they  cohabit  with  other  indi- 
viduals. 

Motility:  After  clinical  observation  of  the  motility  of  the  sper- 
matozoa of  the  bull,  I  find  that  it  differs  little  or  none  from  the 
types  as  observed  by  Eeynolds  (34)  in  his  work  on  human  sperma- 
tozoa. His  ol)servations  are  so  accurate  and  well  described  that 
they  will  be  given  in  his  words.  "All  normal  motions  appear  to 
be  consecutive  phases.  Initial  motion,  i.  e.,  motion  as  seen  in 
fresh  semen  under  favorable  conditions,  consists  of  a  lashing 
of  the  after  part  of  the  tail  from  ^ido  to  side  which  is  so  lapid 


20 

as  to  constitute  vibration.  It  produces  rapid  forward  motion  in  a 
practically  straight  line,  the  head,  middle  piece,  and  forward  por- 
tion of  the  tail  maintaining  their  position  in  the  line  of  motion 
with  practically  no  swaying  from  side  to  side.  The  action  of  the 
flagellum  is  so  rapid  that  it  is  quite  impossible  to  follow  its  indi- 
vidual movements.  Spermatozoa  swimming  in  this  manner  head 
against  the  current  and  usually  cross  the  field  of  observation  in 
about  five  seconds  in  the  absence  of  currents  or  obstacles.  This 
type  of  motion  will  be  described  throughout  the  paper  as  'pro- 
gi'essive  vibratile'  motion.  Progressive  vibratile  motion  is  nor- 
mally succeeded  after  a  variable  length  of  time  by  what  I  regard, 
as  the  second  phase  of  normal  motion. 

"The  second  normal  motion  differs  from  the  first  not  only  in 
its  character  but  in  markedly  reduced  speed.  The  tail  movement 
alters  to  a  long  stroke  from  side  to  side  and  almost  the  whole 
length  of  the  tail  partakes  in  the  stroke.  This  is,  moreover,  accom- 
panied by  swaying  of  the  head  and  middle  piece  through  an  arc 
which  is  always  considerable  and  may  even  equal  ninety  degrees. 
The  general  outline  of  the  spermatozoa,  from  being  practically 
straight  with  almost  non-detectable  sharp,  quick,  small  arc  vibra- 
tion of  the  aftertail,  has  become  an  S  in  outline,  with  large,  slow, 
plainly  perceptible  undulations  traveling  gradually  backward 
throughout  the  length  of  the  spermatozoon.  Speed  has  been  lost 
and  direction  seems  to  be  more  specifically  determined  by  the 
surroundings.  Individuals  at  this  stage  show  a  pronounced  choice 
of  direction  and  go  up  to  objects  in  the  medium,  from  which  they 
later  make  off  as  though  the  movement  were  determined  by  tactile 
reaction  to  some  extent.  This  type  of  motion  has,  therefore,  been 
named  'undulatory  tactile'  in  contradistinction  to  'progressive 
inbratile. ' 

'The  third  type  of  normal  motion  succeeds  the  second  and  con- 
sist.s  in  a  tendency  on  the  part  of  the  spermatozoon  to  push  itself 
against  or  into  any  small  masses  of  cells,  or  sometimes  other 
materials,  which  it  may  find  in  the  neighl^orhood,  bunting  itself 
into  any  small  cove  that  can  be  found,  and  maintaining  a  slight 
burrowing  motion  by  a  lashing  tail  movement  of  the  vibratile  type 
not  unlike  the  movements  of  the  caudal  fin  of  a  fish.  The  move- 
ment of  the  flagellum  in  this  third  type  is  unlike  the  second  type 
in  that  it  is  wlbratile  rather  than  lashing,  but  is  slower  than  the 
\nbratile  motion  of  the  first  type  and  less  limited  to  the  after- 
part  of  the  tail.  The?e  spermatozoa  are  apparently  not  caught 
in  the  debris  or  unable  to  move  off.  From  time  to  time,  they 
back  out  of  such  a  cove  and  seek  another  mooring  place. 

"This  'stationary  bunting'  motion  is  less  universal  than  the 
other  two.  Many  individual  spermatozoa  fail  to  attain  it.  It 
seems  probable  that  only  the  most  vigorous  individuals  ever  reach 
this  stage.  It  has  not  been  encountered  in  unmixed  semen  or  in 
any  artificial  mediums.  It  has  been  observed  only  when  the  sper- 
matozoon is  in  the  secretions  of  the  female  genital  tract.     It  is 


21 

most  frequent  Avhen  the  spermatozoon  is  in  contact  with  a  nest  of 
epithelial  cells     .... 

"The  three  types  of  normal  motion  are  not  only  distinctive  but 
are  always  consecutive,  i.  e.,  the  second  follows  the  first  after 
a  period  which  is  apparently  determined  both  by  initial  vitality 
and  by  the  favorable  or  unfavorable  character  of  the  medium, 
while  the  third  has  been  observed  to  occur  only  in  individuals  which 
have  already  developed  the  second.  They  apparently  constitute  a 
normal  cycle. 

"This  cycle  is  open  to  the  theoretical  explanation  that  these 
types  of  motion  are  directly-  adapted  to  the  function  of  the  sper- 
matozoon ;  thus,  the  progressive  vibratile  motion  which  is  char- 
acteristic of  the  earliest  period  of  its  existence  appears  especially 
suitable  for  its  prolonged  journey  through  the  cervix  and  uterus 
to  the  fundus  and  tube.  This  is  supported  by  the  fact  that  during 
this  motion  it  always  heads  directly  against  any  existing  current, 
and  that  during  this  stage  of  its  journey  it  must  under  natural 
conditions  be  continuously  exposed  to  the  faint  outward  ciliary 
currents  of  the  mucous  membrane  of  these  passages. 

"The  undulatory  tactile  motion  which  succeeds  the  progressive 
vibratile  would  then  be  well  adapted  to  the  later  stage  in  which 
the  spermatozoon  has  reached  the  tube,  where  its  success  in  conju- 
gation is  dependent  on  its  finding  the  ovum  rather  than  on  further 
progress. 

"The  stationary-  bunting  type  of  motion  is  that  which  would  be 
demanded  by  the  passage  of  the  spermat07X)on  through  the  egg 
membrane  which  has  been  so  often  observed  in  the  lower  animals. 
This  very  plausible  explanation  is,  however,  necessarily  theoretical 
and  must  always  remain  so,  as  the  conditions  which  surround  the 
specimen  on  the  field  of  the  microscope  vary  in  so  many  respects 
from  those  in  which  it  accomplishes  conjugation  in  the  course  of 
nature;  but  the  practical  importance  of  the  study  of  tj^pes  of 
motion  is  not  affected  by  their  explanation." 

The  duration  of  motility  is  a  variable  factor,  dependent  entirely 
upon  the  environment  in  which  the  spermatozoa  are  placed. 
Within  the  body  they  usually  sur\'ive  at  least  a  week.  One  author 
describes  a  case  in  which  he  found  living  sperms  in  a  Avoman  who 
stated  that  coition  had  not  been  experienced  for  three  and  one- 
half  weeks.  It  has  been  stated  with  regard  to  human  sperms,  that 
their  motion  should  continue  or  be  cai)able  of  being  re-established 
for  twelve  hours.  Gary  (35)  states,  "first,  that  in  their  proper 
medium  and  at  the  body  temperature  the  viability  of  the  sperm 
cells  may  extend  over  a  period  of  a  few  days ;  second,  that  their 
prolonged  vitality  is  probably  dependent  upon  the  normal  lime 
salts  of  the  prostatic  fluid ;  third,  that  the  sustaining  power  of 
the  seminal  fluid  is  increased  by  its  undon  with  the  normal  secre- 
tion of  the  female  genital  tract."  After  death  of  the  male  animal 
they  retain  their  motilitj'  in  the  genital  tract  for  twenty-four  to 
forty-eight  hours. 


22 

Wolf  (36)  worked  on  this  problem  in  rabbits,  and  summarizes 
as  follows:  "The  motility  of  rabbit  spermatozoa  can  be  preserved 
for  at  least  nine  days  by  placing  the  juice  of  the  epididymis  in 
the  Tyrodc  solution  which  had  been  buffered  and  to  which  glucose 
had  been  added.  The  solution  is  adjusted  to  a  pH  of  about  7.4. 
Oxj'gen  is  passed  in  and  a  suitable  amount  of  sodium  bicarbonate 
added.  The  preparation  must  be  kept  at  a  temperature  near  the 
freezing  i^oint  of  water."  Under  ordinary  conditions  motility 
persists  outside  the  body  only  a  few  hours  after  ejaculation,  but 
if  the  semen  is  kept  quite  cool  till  the  time  of  examination  on  a 
warm  stage,  motility  should  be  capable  of  being  restored  in  at  least 
a  fair  percentage  of  the  sperms  for  twenty-four  to  forty-eight 
hours.  The  cells  are  more  sensitive  to  heat  than  to  cold,  and  even 
to  dilute  acids  more  than  to  alkalies. 

Henle  (quoted  by  Ellenberger)  states  that  a  spermatozoon  under 
favorable  conditions  travels  at  the  rate  of  twenty-seven  millimeters 
in  seven  and  one-half  minutes,  which  makes  three  and  five-tenths 
millimeters  per  minute.  This  is  about  sixty  times  the  entire 
length  of  the  spermatozoon,  and  twenty-one  centimeters  in  an 
hour.  Forward  motion  is  also  more  pronounced  when  the  swim  is 
against  the  current,  such  as  is  produced  by  the  cilia  of  the  oviduct. 
It  has  been  demonstrated  that  feebly  motile  sperms  become  very 
actively  motile  when  placed  on  the  mucosa  of  a  fresh  Fallopian 
tube. 

Technique 

The  material  used  in  the  work  came  from  abattoir  animals,  bull 
calves  and  adult  bulls  raised  in  the  department  herd,  and  sires 
upon  which  clinical  observations  had  been  made  by  various  veteri- 
narians in  the  field.  Senlen  samples,  many  of  which  were  sent  in, 
were  collected  as  often  as  possible  after  the  method  described  by 
AVilliams  (16).  The  genital  organs  were  removed  with  as  little 
chance  of  contamination  as  possible,  and  taken  or  sent  to  the 
la1)oratory  where  the  examinations  were  made  soon  after  arrival. 

All  cultures  were  made  by  searing  the  surface  carefully,  tearing 
out  a  small  portion  of  the  tissue  with  sterile  forceps,  and  placing 
it  upon  the  media.  In  most  cases,  however,  where  fluids  were 
])resent,  tubes  were  inoculated  with  the  material  which  had  been 
drawn  off  with  a  sterile  pipette.  As  stated  by  Carpenter  (9),  in 
his  work  on  the  female  genital  tract,  the  organisms  usually  live 
in  the  depths  of  the  tissue.  The  media  used  principally  were  glu- 
cose glycerin  agar  (glucose  1  per  cent,  glycerin  3  per  cent)  ;  plain 
agar,  both  with  a  pH  value  of  7.4,  and  Loeffler's  blood  serum. 
Small  amounts  of  sterile  blood  serum  or  defibrinated  blood  were 
added  to  most  of  the  agar  slants  to  insure  better  growths  of 
streptococci  when  present.  All  tubes,  to  which  the  serum  had 
been  added,  were  incubated  for  forty-eight  hours  before  inocula- 
tion to  insure  absolute  sterility. 

After  inoculation,  the  agar  tubes  were  sealed  with  sealing  wax 
to  give  a  partial  oxygen  tension  which  was  quite  necessary  in  iso- 


23 

lating  the  streptococci.  The  growth  of  other  organisms  was  by 
no  means  hindered  by  the  procedure,  for  one  tube  from  eacii 
organ  was  often  left  unseaKxl.  Incubation  was  at  ol'^  C,  and  the 
routine  method  of  examining  the  tubes  was  identical  with  the 
method  of  Carpenter  (9). 

Whenever  possible,  a  sample  of  blood  was  obtained  from  the 
animal  either  before,  or  at  the  time  of  slaughter,  for  agglutination 
wuh  Bact.  aburtum  antigen.  Extracts  from  the  semnial  vesicles, 
testes,  and  epididymes  were  injected  into  the  guinea  pigs  and 
examined  at  the  end  of  four  to  six  weeks  for  the  presence  of 
Bad.  ahortum. 

Sections  of  all  organs  were  fixed  as  soon  as  possible  in  either 
Zenker's  or  Ilelly's  fluid.  Hematoxylin  and  eosin  were  used  as 
routine  tissue  stains.  Eosin  and  methylene  blue,  and  Mallory's 
connective  tissue  stain  Avere,  however,  frequenth'  utilized  for 
special  staining  reactions. 

The  motility  of  the  spermatozoa  is  best  observed  about  half  an 
hour  after  ejaculation,  when  the  thick  tenacious  clot  has  started 
to  liquefy.  A  drop  of  the  fluid  is  placed  upon  a  warmed  slide, 
preferably  one  with  a  slight  depression  in  it.  and  observation 
made  with  high  or  low  powered  objectives.  The  semen  may  be 
examined  whole,  or  diluted  with  physiological  saline  .solution.  In 
the  latter  ease,  the  sperms  have  a  greater  opportunity  for  freedom 
of  motion  in  the  absence  of  the  thick  viscid  coagulate.  A  small 
vial  erf  saline  solution  may  be  carried  in  one's  pocket  where  it 
will  be  kept  warm,  and  a  drop  of  this  placed  upon  the  glass  slide. 
If  the  clot  of  semen  is  merely  touched  to  this  drop  on  the  slide, 
plenty  of  .spermatozoa  will  be  deposited  for  an  examination.  This 
method  is  very  satisfactory  for  the  observation  of  motility,  but 
needless  to  say,  the  undiluted  semen  must  be  used  for  the  deter- 
mination of  the  number  of  sperms  present.  If  necessary,  the 
specimen  may  be  covered  with  a  cover  glass  and  the  oil  immersion 
objective  used.  While  a  warmed  slide  is  quite  sufficient  to  enable 
one  to  detect  the  i^resenee  of  motility,  the  field  soon  cools  and  the 
sperms  gradually  become  less  motile.  If  possi])le.  it  is  best  to 
use  a  small  electrically  heated  stage  warmer,  which  keeps  the 
field  at  a  constant  ])ody  temperature,  so  that  the  duration  of  the 
motion  may  be  observed  for  hours  if  warmed  jihysiological  saline 
solution  is  added  as  the  fluid  evaporates. 

Stained  preparations  are  best  made  with  fhin  smears  on  the 
glass  slides.  This  is  conveniently  done  by  placing  a  drop  of  the 
semen  on  a  slide  and  smearing  it  over  the  surface  with  the  edge 
of  another  slide.  A  fairly  thin  and  even  field  is  thus  obtained.  A 
.still  better  method  is  to  first  dilute  the  semen  with  physiological 
saline  solution,  so  as  to  ol)tain  fewer  sperms  in  the  field.  After 
drying  the  pre]>aration  in  air.  fixation  may  l)e  produced  by  draw- 
ing the  slide  through  a  gas  flame  several  times,  by  immersion  in 
equal  parts  of  alcohol  and  ether,  or  even  1)y  the  use  of  tissue  fixers 
such  as  Ilellv's  or  Zenker's  .fluids.      For  ordinary  .staining,  heat 


24 

fixation  is  the  quickest,  and  at  the  same  time  is  quite  satisfactory. 
After  the  slide  is  cooled,  or  washed,  to  remove  the  fixing  solutions, 
it  should  be  placed  for  a  few  minutes  in  a  freshly  prepared  solu- 
tion (1  per  cent)  of  chlora/^ene,  as  recommended  by  Williams,  to 
remove  the  mucus  and  proteid  material  which  otherwise  blur  the 
field.  Other  authors  (38)  have  recommended  diluting  the  semen 
with  about  twenty  volumes  of  a  0.12  per  cent  sodium  carbonate 
solution  in  0.8  per  cent  sodium  chloride.  From  this  liquid  the 
cells  should  be  centrifuged  for  several  minutes,  removed  with  a 
pipette,  and  smeared  on  the  slide.  Following  this,  the  slide  should 
be  thoroughly  washed,  preferably  for  ten  minutes  in  running 
water,  after  which  it  is  ready  for  staining.  Numerous  methods 
have  been  used  for  this  procedure,  but  the  sperms  are  more  or 
less  erratic  in  their  reactions  to  the  dyes,  and  one  must  be  very 
careful  to  use  the  same  method  in  all  samples,  in  order  to  obtain 
uniform  results.  For  quick  staining,  to  bring  out  gross  abnormal- 
ities of  structure,  and  number  of  sperms  present,  one  may  use 
Gram's  stain,  or  a  light  stain  with  any  of  the  aniline  dyes,  such 
a.s  fuchsin.  To  bring  out  the  finer  structure,  particularly  of  the 
head,  more  careful  technic  must  be  employed. 

Carnett  and  others  (38)  recommend  the  following:  "  The 
method  of  staining  by  iron-hematoxylin,  particularly  when  supple- 
mented by  a  cytoplasm  stain,  has  proved,  on  the  whole,  the  most 
satisfactory,  and  possesses  the  additional  advantage  of  being  abso- 
lutely permanent,  a  quality  that  few  anilines  can  boast.  The 
method  consisted  of  treating  the  fixed  object  —  and  here  the  fixing 
agent  was  heat  —  with  a  two  per  cent  solution  of  iron-alum  for 
from  two  to  four  hours.  The  excess  of  iron-alum  was  then  com- 
pletely removed  1)y  pure  water,  and  the  object  treated  with  a  solu- 
tion of  hematoxylin  (one  per  cent  aqueous)  for  twelve  hours  or 
longer.  The  cells  by  this  time  were  perfectly  black.  However,  a 
1  per  cent  solution  of  iron-alum  removed  the  stain  from  the  cyto- 
pla.sm.  leaving  the  chromatin  of  the  head,  the  centrosome,  and  the 
axial  filament  a  brilliant  blue-black.  Care  must  be  taken  that  the 
preparation  is  not  over-decolorized.  After  deeolorization  a  satu- 
rated aqueous  solution  of  eosin  was  added  for  from  one  to  three 
minutes.  This  stained  the  protoplasmic  envelope  pink,  and,  unless 
the  envelope  is  overstained,  the  view  of  the  inner  structures  is  not 
impaired  in  the  least." 

AVilliams  (17)  recommends  using  two  staining  solutions,  one  of 
alcoholic  eosin  and  fuchsin,  the  other  a  diluted  methylene  blue. 
The  results  obtained  are,  however,  more  or  less  erratic,  due  to 
the  unstable  character  of  the  former  stain,  and  the  ease  with 
which  one  may  over  or  under  stain.  Many  beautiful  specimens 
may,  nevertheless,  be  obtained  by  this  method.  I  have  frequently 
used  a  fairly  quick  method,  though  one  not  satisfactory  in  ail 
cases,  which  consists  of  staining  for  five  or  six  minutes  in  a  satu- 
rated aqueous  solution  of  fuchsin,  washing  in  water,  and  counter- 
staining  for  a  few  seconds  in  a  strong  solution  of  methylene  blue. 


25 

A  quite  satisfactory  method  is  to  stain  from  two  to  five  minutes  in 
a  saturated  aqueous  solution  of  methyl  green,  with  the  application 
of  gentle  heat.  The  heat  may  be  applied  by  warming  the  slide  over 
a  gas  tiame  as  it  steams,  or  by  placing  the  jar  containing  the  stain 
in  a  hot  water  bath.  The  slide  is  then  washed  thoroughly  and 
eounterstained  for  five  minutes  in  a  strong  aqueous  solution  of 
eosin.  This  is  a  fairly  reliable  method,  and  many  excellent  pre- 
parations may  be  obtained  by  its  use.  The  nucleus  is  stained 
green,  the  anterior  part  of  the  head  and  all  of  the  tail  pink.  So 
far,  I  have  found  this  a  very  reliable  stain  for  routine  work. 

Pathology 

In  the  genital  tracts  that  I  have  studied,  a  complete  pathological 
and  bacteriological  examination  was  made  Avherever  possible,  but 
in  man}'  of  the  abattoir  animals,  and  certain  others,  gross  and 
microscopical  examinations  only  could  be  made.  The  genital 
organs  of  one  hundred  and  ninety-six  males  have  been  examined, 
and  the  gross  or  miscroseopical  changes,  or  both,  determined.  The 
abattoir  animals  Avere  from  a  large  slaughter  house,  and  a  small 
local  plant. 

Of  the  tracts,  the  pathology  of  which  was  studied,  two  were 
from  aborted  fetuses ;  seven  from  apparently  normal  young  calves ; 
four  from  mature  fertile  bulls ;  and  sixteen  from  mature  infertile 
or  sterile  sires.  The  remainder  (167)  of  those  examined  were 
from  abattoir  animals.  Besides  these,  three  specimens  of  seminal 
vesicles,  and  seven  of  testes  were  studied. 

The  tracts  of  the  aborted  fetuses  and  veal  calves  were  apparently 
normal,  both  on  gross  and  microscopic  examination  of  the  vesicles 
and  testes.  On  gross  examination,  the  tracts  of  the  mature  fertile 
bulls  were  normal,  except  for  the  presence  of  many  fine  connective 
tissue  tufts  and  strands  upon  the  serous  covering  of  the  tails  of 
the  epidid>Tnes,  and  adjacent  portion  of  the  parietal  layer  of  the 
tunica  vaginalis.  jMicroscopic  sections  of  all  parts  were  apparently 
normal.  The  more  important  pathological  changes  in  the  tracts  of 
the  sixteen  sterile  or  infertile  bulls  are  given  in  the  appended 
chart.  The  tracts  are  numbered  the  same  as  in  Group  VI  of  the 
report  of  the  bacteriological  findings ;  that  is,  any  particular  num- 
ber in  either  tal)le  refei's  to  the  same  animal.  Eeferences  are  made 
throughout  the  text  to  some  cases  which  appear  in  this  group  of 
animals.  Prostate  and  Cowper's  glands  are  not  included  in  the 
chart  as  they  were  not  examined  in  some,  and  were  negative  in  the 
others.  Fibrous  tufts  and  strands  were  present  on  the  covering 
of  the  epididymes  in  each  animal. 

The  study  of  sections  from  the  abattoir  animals,  as  well  as  those 
from  the  sterile  or  infertile  bulls,  forms  the  basis  for  the  follow- 
ing observations  upon  the  pathologA'  of  the  male  genital  tract. 
The  tracts  secured  from  the  abattoir  were  studied  for  the  most 
part  on  the  ba.sis  of  the  organ  rather  than  on  that  of  the  animal. 
For  example,  all  sections  of  testes  were  placed  in  the  same  bottle 
of  fixer,  and  the  same  plan  followed  for  the  other  organs. 


26 

Testes  :  The  testes  seldom  presented  gross  alterations  of  struc- 
ture except  for  abscess  formation,  which,  according  to  Williams, 
occurs  more  frequently  in  the  bull  than  in  any  of  the  other 
domesticated  animals.  He  also  states  that  arrest  in  development  by 
which  the  organs  remain  soft,  flaccid,  and  somewhat  smaller  than 
normal  is  not  uncommon.  One  very  interesting  specimen,  which 
typifies  abscess  formation,  came  from  a  bull  with  a  history  reveal- 
ing that  one  testis  had  become  much  enlarged,  hot,  and  painful. 
These  sj^mptoms  developed  very  rapidly.  Anorexia  was  well 
marked.  Local  applications  were  used  for  several  weeks,  but 
at  the  end  of  two  months  the  condition  was  so  little  improved  that 
unilateral  castration  Avas  performed.  The  general  condition  of  the 
animal  soon  improved,  but  after  a  year  of  service  he  was  so  uncer- 
tain compared  to  what  he  had  been,  that  he  was  sent  to  the 
butcher.  It  was  impossible  to  obtain  the  other  testicle  for  study, 
though  it  undoubtedly  was  abnormal.  The  testicle  removed  was 
considerably  enlarged,  measuring  twenty  by  ten  and  one-half  centi- 
meters. The  tunica  albuginea  presented  a  thickness  of  six  milli- 
meters, and  was  made  up  of  firm  sclerotic  tissue.  The  epididy- 
mis was  not  recognizable  in  the  mass.  Testicular  tissue  was  almost 
entirely  gone.  The  only  remains,  of  what  appeared  to  have  been 
I)arenchyma,  was  an  elongated  irregular  area  at  one  side  of  the 
organ.  This  tissue  consisted  of  a  whitish  opalescent  material, 
speckled  with  varying  sized  abscesses.  This  organ  is  pictured  in 
Fig.  3.  The  remainder  of  the  organ  consisted  of  a  thick  yellowish 
caseous  mass.  Strepiococcns  viridans  was  recovered  from  the 
outer  portion  of  the  organ,  and  guinea  pig  inoculations  failed  to 
demonstrate  Bact.  ahortum. 

^Microscopically,  changes  are  quite  common  and  varied  in  char- 
acter. In  the  seminiferous  tubules,  the  changes  range  from  a 
slight  desquamation  of  the  germinal  epithelium  to  atrophy  and 
complete  degeneration  of  the  entire  tubule,  as  was  the  case  in  the 
left  testis  of  Bull  1.  In  the  mild  cases,  spermatogenesis  occurs 
apparently  in  a  normal  manner  up  to  the  spermatid  stage,  at 
wliich  point  many  of  the  cells  degenerate  and  slough  off.  These 
appear  in  the  seminal  fluid,  associated  with  the  few  sperms  that 
reach  maturity.  This  sloughing  and  degeneration  may  be  localized 
in  a  few  of  the  tubules,  or  it  may  be  widespread  over  the  entire 
organ.  Likewise,  the  changes  may  involve  not  only  the  more 
mature  cells,  but  they  may  be  so  .severe  as  to  cause  almost  total 
degeneration  and  desquamation  of  the  seminal  epithelium,  as  in 
Fig.  15.  These  defects  in  spermatogenesis  are  of  course  evidenced 
in  the  semen  l)y  the  presence  of  immature,  or  abnormal  types  of 
sperms.  With  cessation  of  spermatogenesis  or  degeneration  of  the 
epithelium  of  the  entire  gland,  no  sperms  are  formed.  Not  infre- 
(inently  one  finds  numerous  tubules,  or  even  the  entire  testis  in 
which  the  germinal  epithelium  is  intact,  ])ut  there  is  little  or  no 
evidence  of  mitosis,  as  in  some  tubules  of  Bull  6.  The  cells  are 
several  layers  deep  as  in  the  normal  condition,  but  they  are  not 
dividing.    This  condition  is  shown  in  Fig.  13. 


27 

In  the  more  chronic  forms,  the  tubules  become  atrophied,  and 
frequently  disappear  entirely.  The  membrana  propria  may  be- 
come thickened,  due  to  excessive  connective  tissue  formation,  or 
infiltration  with  serum  or  exudate.  On  the  other  hand,  a  dis- 
tinct atrophy  may  occur.  The  sti'oma  of  the  organ  not  i)ifre- 
([uently  is  thickened  by  inflammatory  exudates,  or  by  a  noticeable 
increase  in  the  connective  tissue.  In  some  testes,  the  connective 
tissue  is  so  much  increased  that  the  tubules  rapidly  become 
atrophied,  and  disappear.  In  abscess  formation,  due  to  acute  in- 
flammations, the  entire  organ  Ijecomes  enlarged,  markedly  hyper- 
aemic,  and  infiltrated  with  leucocytes.  Necrotic  areas  appear  here 
and  there  in  the  parenchyma.  The  rete  often  shows  a  marked  de- 
generation of  the  lining  epithelium,  and  atrophy  caused  by  in- 
crease of  the  interstitial  connective  tissue. 

Epididymis:  This  organ  not  infrequently  presents  gross  ab- 
normalities, and  very  often  is  pathological  on  microscopic  examina- 
tion. Acute  inflammation,  with  induration  or  abscess  formation,  is 
very  common  in  the  tail,  but  less  so  in  the  head  and  body. 
Possibly  this  is  caused  by  the  fact  that  the  tail  is  the  most  pendant 
portion  of  the  organ.  In  these  cases,  the  tail  is  enlarged,  soft,  and 
quite  hot  and  painful  on  physical  examination.  Enlargement,  due 
to  a  connective  tissue  induration,  occurs  occasionally  in  all  three 
parts,  and  the  inflammation  may  produce  adhesions  to  the  adjacent 
serous  membranes.  Inflammation  of  both  the  parietal  and  visceral 
layers  of  the  tunica  vaginalis  is  very  common.  In  those  cases, 
the  membrane  usually  is  quite  hyperaemic.  and  on  the  surface  it 
presents  many  small  reddened  tufts  of  newly-formed  connective  tis- 
sue. In  adult  bulls  it  is  exceptional  not  to  find  at  least  slight  evidence 
of  some  previous  inflammation.  In  all  of  the  numerous  bulls  ex- 
amined, both  apparently  normal  and  sterile,  I  have  found  but  one 
in  which  some  evidence  of  inflammation  (either  present  or  past) 
could  not  be  found.  Along  with  the  fibrous  tufts,  are  numerous 
fine  strands  of  connective  tissues  passing  from  the  covering  of  the 
tail  of  the  epididymis  to  the  adjacent  portion  of  the  parietal  layer 
of  the  tunica.  The  strands  often  extend  even  to  the  upper  part  of 
the  head. 

INIicroscopically,  inflammation  of  the  part  is  shown  by  hyper- 
aemia,  loss  of  cilia  of  the  lining  cells,  and  exudation.  In  the  more 
.severe  forms,  the  lining  cells  which  furnish  considerable  secretion 
for  the  nourishment  and  stimulatioji  of  the  sperms,  l)ecome  de- 
generated, and  are  exfoliated  into  the  lumen,  as  in  Fig.  21.  This 
condition  is  very  common  in  sterile  bulls,  and  those  of  lowered 
fertility.  In  the  chronic  types,  the  interstitial  connective  tissue  is 
increased  in  amount,  leading  to  degeneration  and  atrophy  of  part 
or  all.  of  the  tubules,  as  in  the  case  of  Bull  2.  Infiltration  with 
leucocytes,  and  necrosis,  are  the  predominating  lesions  in  the  p.vo- 
genic  types  of  inflammation. 


28 

Ductus  Deferens  :  This  tube  seems  to  be  peculiarly  free  from 
severe  inflammatory  processes,  and  when  these  appear  they  are 
limited  to  the  mucosa.  The  cells  of  the  lining  membrane  not  in- 
frequently  show  a  mild  type  of  degeneration  and  exfoliation,  or  in 
the  more  chronic  forms,  the  entire  membrane  degenerates  and  disap- 
pears.  In  man,  the  duct  occasionally  becomes  occluded,  but  so  far 
I  have  failed  to  find  this  condition  in  the  bull.  Undoubtedly,  when 
the  occlusion  does  occur,  it  is  near  the  origin  of  the  duct  at  the 
tail  of  the  epididymis. 

Seminal  Vesicles:  The  seminal  vesicles  and  epididymis, 
especially  the  tail,  seem  to  be  the  parts  most  subject  to  extensive 
pathological  changes,  and  bacterial  invasion.  In  most  instances, 
diseased  vesicles  present  gross  manifestations  recognizable  on 
clinical  examination,  while  on  the  other  hand  microscopic  changes 
may  be  present  in  the  absence  of  gross  lesions.  As  diagnosed  on 
physical  examination,  or  even  on  post  mortem  examination  of  the 
tract,  the  various  forms  may  be  classified  into : 

1.  Acute  Catarrlial  Type:  In  this  form,  the  vesicles  are  usually 
enlarged,  soft,  and  more  or  less  reddened  by  hyperaemia.  On 
physical  examination  of  the  animal,  distinct  flinching  is  produced 
when  pressure  is  applied  to  the  organ.  Enlargement  may  even 
be  absent  in  the  early  stages,  and  the  diagnosis  may  be  made  from 
the  extrenie  sensitiveness  alone. 

2.  Suppurative  or  Cystic  Types:  In  both  of  these  types,  the 
vesicles  are  usually  enlarged,  either  uniformly,  or,  as  is  usually  the 
case,  in  localized  areas.  The  suppurative  form  may  extend  over 
the  entire  gland,  forming  one  large  encapsulated  abscess,  or  on  the 
other  hand,  it  may  take  the  form  of  variable  sized  abscesses  with 
thick  sclerotic,  or  thin  fluctuating  walls.  Occasionally  the  abscesses 
rupture  and  discharge  their  contents  into  the  rectum.  Dr.  Wil- 
liams presented  one  case  of  this  type.  One  vesicle  was  apparently 
normal,  whereas  the  other  was  about  five  times  larger  than  normal, 
and  consisted  of  a  dense  outer  capsule  which  was  adherent  to  all 
surrounding  parts.  On  dissection,  it  was  found  that  the  organ 
consisted  of  abscesses  of  various  sizes,  the  larger  one  of  which  had 
ruptured  some  time  previously  into  the  rectum,  leaving  the  dis- 
tinct remains  of  an  opening  into  that  part.  The  cystic  form  may 
occur  either  with  or  without  suppuration.  One  case  came  to  my 
attention  in  which  both  vesicles  were  made  up  of  abscesses  of 
varying  sizes  as  well  as  of  a  smaller  number  of  cysts.  Evidently 
the  cysts  were  of  the  retention  type,  and  were  secondary  to  the 
pyogenic  infection, 

3.  Chronic  or  Sclerotic  Type:  This  form  is  characterized  by  a 
distinct  firmness  with  or  without  marked  enlargement.  The  con- 
dition may  be  accompanied  by  disease  of  the  parenchymatous  tissue 
or  it  may  take  the  form  of  a  chronic  productive  inflammation  of 
the  interstitial  tissue.  This  inflammation  may  be  simply  a  super- 
ficial thickening,  or  it  may  extend  in  between  the  lobules. 


29 

4.  The  Peri-vesicular  or  ^'pan-inflammatory"  Type  usually  is 
the  result  of  severe  inflammation  of  the  vesicles,  with  probable  rup- 
ture of  some  of  the  smaller  cysts  or  abscesses  upon  the  surface. 
The  vesicles  are,  as  a  rule,  considerably  enlarged  and  buried  in  a 
dense  mass  of  adhesions  which  involve  neighboring  structures. 
The  vesicles  cannot  be  palpated  on  i)hysical  examination,  and  it  is 
only  on  careful  post  mortem  dissection  that  they  may  be  studied. 
This  type,  however,  is  quite  rare, —  two  cases  only  having  come  to 
my  attention.  In  both,  the  vesicles  themselves  were  markedly 
affected. 

Microscopically,  changes  in  the  vesicles  are  quite  frequently  en- 
countered, even  in  the  absence  of  gross  manifestations.  In  the 
•acute  catarrhal  forms,  the  mucosa  and  submucosa  are  hyperaemie. 
The  lining  cells  show  various  forms  of  degeneration,  and  there 
are,  as  a  rule,  inflammatory  exudates  in  the  lumen.  As  the  in- 
flammation progresses,  the  lining  cells  degenerate  further,  and 
become  cast  off  into  the  lumen  of  the  glandular  cavities,  as  in 
Plate  VI.  The  normal  clear  mucous  secretion  becomes  mixed  with 
fibrin,  leucocytes,  and  cellular  debris.  These  changes  may  involve 
merely  parts  of  the  organ,  or  they  may  be  quite  extensive.  With 
large  sections,  one  may  find  the  inflammation  in  all  stages,  from  the 
mildest  catarrhal  type,  to  complete  degeneration  and  exfoliation 
of  the  secretion-forming  mucosal  cells,  and  filling  of  the  cavities 
with  degenerated  cells,  leucocytes,  and  debris.  Frequently  the  inter- 
stitial tissue  is  in  no  way  affected,  but  at  times  it  is  thickened  by 
©edematous  exudates,  leucocjiies,  and  fibrin.  The  chronic  inter- 
stitial form  is  characterized  by  a  considerable  increase  of  con- 
nective tissue, —  producing  marked  atrophj',  or  even  complete 
obliteration  of  the  glandular  cavities.  Microscopically  the  sup- 
purative form  may  be  diffuse  over  the  entire  gland,  or  as  stated 
previously,  may  be  in  the  form  of  localized  abscesses,  with  or  with- 
out a  thick  connective  tissue  Avail.  The  parenchyma  in  these  cases 
is  usually  extensively  degenerated  and  atrophied  in  those  parts 
that  have  not  undergone  suppuration  and  necrosis.  The  cysts 
appear  to  be  of  the  ordinary  retention  type,  and  may  or  may  not 
be  accompanied  by  extensive  changes  in  the  lining  epithelium. 

Both  the  abscess  formation  and  cystic  conditions  are  undoubt- 
edly initiated  by  an  obstructive  inflammation  of  all  or  part  of  the 
excretory  duct.  This  is,  however,  a  protective  mechanism,  for 
Avhere  the  duct  is  closed  the  bacteria  and  exudates  are  unable  to 
reach  the  urethra  and  contaminate  the  semen. 

Prostate  and  Cowper's  Glands:  These  glands  were  more  or 
less  neglected  in  the  early  part  of  the  work,  but  later  were  sub- 
jected to  the  same  examination  as  other  parts.  Of  the  thirty-six 
of  each  type  of  gland  examined,  I  failed  to  find  one  with  any  gross 
changes,  l)ut  two  prostates  were  found  that  presented  a  mild 
catarrhal  inflammation  of  the  mucosa.  It  is  probable  that  Cow- 
per's  glands,  as  well,  occasionally  undergo  inflammatory  changes. 


30 

Semen  :  The  semen,  made  up  as  it  is  of  mixed  products  of 
the  testes  and  accessory  sexual  glands,  is  very  often  abnormal,  as 
would  be  expected  in  view  of  the  frequency  with  which  changes 
occur  in  the  glands  contributing  to  its  formation.  The  normal 
semen  is  remarkably  adapted  to  its  function  of  nourishment  and 
stimulation  of  the  sjiermatozoa,  and  their  conveyance  to  the  in- 
ternal female  genital  organs.  The  spermatozoa  are  extremely 
sensitive  to  changes  in  their  environment,  with  the  result  that  any 
alteration  of  the  physical  or  biochemical  content  of  the  seminal 
fluid  may  cause  death  of  the  sperms.  With  this  in  view,  we  must 
remember  that  disease  of  any  of  the  contributing  organs  is  a  poten- 
tial danger,  and  threatens  the  potency  of  the  animal.  Each  or  all 
of  the  glands  may  add  bacteria,  acid  secretions,  or  inflammatory 
exudates.  On  the  other  hand,  they  may  not  function  at  all.  In 
each  case,  however,  the  semen  is  altered. 

Unfortunately  it  is  impossible  with  present  methods  to  obtain 
the  fluid  absolutely  free  from  vaginal  mucus,  but  with  care  it  may 
be  secured  reasonably  free  from  contamination  by  douching  the 
prepuce  of  the  bull  and  vagina  of  the  cow  before  service.  This 
method  was  used  as  often  as  possible  in  collecting  the  samples. 
The  usual  amount  of  semen  obtained  was  from  six  to  ten  cubic 
centimeters. 

With  a  hj'persecretion  of  one  or  all  of  the  glands,  the  semen 
becomes  quite  thin  and  watery,  with  a  deficiency  of  solid  matter, 
together  wdth  changes  in  reaction.  On  the  other  hand,  hypofunc- 
tion  results  in  a  secretion  too  viscid,  which  is  equally  unsuited  to 
the  requirements  of  the  spermatozoa.  The  thin  watery  semen 
clots  imperfectly  or  not  at  all,  and  clotting  is  essential  in  protect- 
ing the  spermatozoa  from  the  acid  secretions  of  the  vagina. 
Likewise,  a  medium  too  viscid  is  a  distinct  hindrance  to  motility. 
Changes  in  reaction  are  very  frequently  encountered.  The  sperms 
are  very  sensitive  to  dilute  acids,  so  that  with  even  a  slight  acidity 
motility  may  diminish  or  entirely  cease.  Purulent  inflammatory 
exudates  are  occa.sionally  mixed  with  the  semen,  and  although  the 
pus  cells  themselves  have  not  been  found  to  be  destructive  to  the 
sperms,  certain  degeneration  products  in  the  exudate  are  very 
toxic,  and  inhibit  or  destroy  the  motility.  So  far,  I  have  failed 
to  find  red  corpuscles  present.  One  very  interesting  sample  of 
semen  was  quite  thick,  of  a  yellowish  green  color,  and  of  a  dis- 
tinctly acid  reaction.  The  secretion  from  the  vesicles  was  later 
found  to  be  of  this  same  character,  and  was  due  to  a  Ps.  pxjocija- 
neus  infection.  The  vesicles  were  highly  inflamed  and  degenerated. 
The  spermatozoa  were  in  this  case  markedly  decreased  in  number, 
and  devoid  of  motility. 

The  early  precipitation  of  the  "Boettcherchen."  crystals  S€ems 
to  be  intimately  connected  with  sterile  semen,  or  spermatozoa  of 
lowered  vitality.  Likewise,  a  decrease  in  solid  matter  is  often  seen 
in  a  deficient  secretion.  In  normal  semen,  the  clot  disappears  after 
standing  a  time,  and  a  thick  .sediment  settles  out.  This  sediment  is 
decreased  in  amount  as  a  rule  in  abnormal  semen. 


31 

Spermatozoa  :  Spermatozoa,  the  essential  germinal  elements, 
are  very  frequently  abnormal,  changes  in  which  may  be  manifested 
in  many  ways.  We  may  divide  the  deviations  into  changes  in 
structure,  and  changes  in  the  motility  which  is  so  indicative 
of  the  intrinsic  vitality  of  the  sperm.  Reynolds  (34)  describes 
two  forms  of  abnormal  motion.  The  first  is  "  rotary  swim- 
ming," in  which  the  sperms  move  forward  progressively,  and 
sometimes  with  fair  rapidity,  but  in  a  spiral  screwlike  manner,  lie 
states  that  this  type  of  swimming  is  very  awkward,  easy  to  recog- 
nize, and  is  usually  of  quite  long  duration.  The  other  form  termed 
"pendulum  swimming,"  he  slates,  is  less  common  than  the  rotary 
swimming  and  is  usually  confined  to  relatively  fewer  sperms  in  a 
given  field.  "In  this  the  middle  piece  and  upper  tail  seem  to  lose 
their  flexibility  and  balance  to  a  considerable  degree,  and  the 
lower  tail  motion  swings  the  forward  part  of  the  spermatozoon  to 
and  fro  with  a  pendulum  movement.  This  type  oi:  swimming  yields 
very  poor  progress." 

One  factor  we  must  bear  in  mind  in  the  study  of  the  semen  ob- 
tained from  the  vagina,  is  that  the  spermatozoa  may  be  highly 
motile  before  ejaculation,  but  the  admixture  of  hostile  vaginal 
mucus  may  inhibit  or  destroy  the  motility.  On  the  other  hand, 
the  conditions  may  be  reversed.  Gary  (35),  in  one  instance, 
found  that  the  spermatozoa  in  a  sample  of  semen  collected  from  a 
condom,  appeared  to  be  of  very  low  vitality,  Avhile  when  thej^  were 
mixed  with  the  vaginal  secretions,  an  (exaggerated  activity  was 
manifested.  INIay  we  not  have  to  contend  with  this  factor  in  some 
herds  in  which  there  is  a  very  distinct 'acid  and  toxic  vaginal 
secretion  from  the  products  of  cervicitis  and  vaginitis'? 

In  a  study  of  motility,  we  must  consider  not  only  the  abnormal 
types  which  may  be  encountered  but  the  percentage  of  motile  cells, 
and  the  duration  of  the  movement.  In  nccrospcrmia  all  the 
ejaculated  cells  are  motionless  or  dead.  In  other  specimens,  vary- 
ing percentages  of  the  cells  are  without  motion,  and  the  others  may 
be  possessed  of  full  and  lasting  motility.  On  the  other  hand,  the 
motility  in  some  eases  is  veiy  active  at  first,  Init  quickly  subsides 
even  undin*  the  best  of  conditions.  The  appearance  in  freshly 
ejaculated  semen  of  numerous  sperms  that  have  a  tendency  to  take 
on  the  "undulatory  tactile"  type  of  motion  when  they  should  be  in 
a  hitxhly  active  state,  is  very  indicative  of  lowered  vitality.  Many 
specimens  present  this  very  picture,  whereas  the  very  active  pro- 
gressive movement  should,  under  proper  conditions,  survive  for  a 
considerable  time  before  it  gives  way  to  the  second,  and  slower 
type.  The  cells  frequently  enrly  bunt  into  epithelial  cells  or  cUimps 
of  immotile  sperms,  then  back  out  and  move  around  sluga'ishly, 
only  to  repeat  the  same  performance  till  they  stop  moving  en- 
tirely. I  have  seen  one  specimen  in  which  the  sperms  all  tended 
to  clump.  Whether  this  was  the  result  of  some  agglutinative  sub- 
stance in  the  vaginal  secretion  is  problematical.  I  have  seen  several 
specimens  of  semen  in  which  practically  all  the  sperms  were  motile 


32 

when  first  examined,  but  the  motion  did  not  sursdve  for  any  great 
length  of  time.  Even  a  small  percentage  of  motionless  sperms  or 
of  those  sho^\^ng  lowered  vitality  is  a  considerable  factor  in 
potency.  xVlthough  millions  of  the  germinal  elements  are  ejacu- 
lated into  the  vagina,  large  numbers  of  them  are  destroyed  or  be- 
come motionless  there,  and  a  small  number  is  left  behind  in  the 
cervix  and  uterus ;  so  that  even  though  but  a  single  sperm  is 
required  for  fertilization,  the  chances  of  impregnation  are  dimin- 
ished in  proportion  to  the  number  of  dead  or  defective  sperms. 

Aspermia:  Absence  of  spermatozoa  in  the  semen  is  rarely  en- 
countered, and  is  probably  due  either  to  total  cessation  of  sperma- 
togenesis, or  to  an  obstruction  at  some  point  in  the  system  of 
excretory  ducts.  I  have  seen  but  one  case  of  this  character.  The 
semen  of  this  bull  was  greatly  increased  in  amount,  and  of  a  thin 
watery  consistency.  Due  to  lack  of  cooperation  on  the  part  of  the 
owners,  the  tract  could  not  be  obtained  for  study.  Oligospermia, 
or  a  diminution  of  the  number  of  spermatozoa,  is  quite  common, 
and  is  undoubtedly  associated  with  defective  spermatogenesis, 
either  as  a  result  of  poor  mitosis  of  the  seminal  epithelium,  or 
degeneration  of  the  elements  before  maturity.  This  condition  may 
vary  from  the  finding  of  only  occasional  dead  sperms  in  the  field,  to 
but  a  slight  decrease  in  the  usual  number  of  normal  sperms 
observed. 

Abnormalities  in  morphology  may  be  classified  into  immature 
types,  and  deformities  or  imperfect  development  of  the  head  and 
tail.  Defective  spermatogenesis  occurs  so  frequently  that  it  is 
not  surprising  to  find  spermatozoa  in  various  stages  of  develop- 
ment cast  into  the  excretory  ducts.  The  various  stages  passed 
through  in  the  development,  from  spermatogonia  to  adult  sperm,  are 
numerous,  and  it  therefore  is  to  be  expected  that  we  should  see  in 
abnormal  semen  many  different  immature  forms.  No  classification 
of  the  various  types  can  be  made,  but  a  clearer  understanding  of 
them  can  best  be  obtained  by  a  review  of  the  process  of  spermato- 
genesis. 

Spermatocytes  and  spermatids  are  seen  in  the  more  severe  types 
of  defective  spermatogenesis,  and  are  relatively  uncommon,  while 
the  more  mature  forms  that  result  from  the  transformations  of 
spermatid  to  adult  cell  are  very  often  seen.  Some  of  these  inter- 
mediate types  are  large  oval  cells  without  distinct  nuclei  and  as  a 
rule  with  poorly  developed  tails.  Cells  with  no  tails  or  distinct 
nuclei,  those  with  protoplasmic  appendages  to  the  head  or  tail,  and 
various  other  types,  are  occasionally  encountered.  IMost  of  these 
are  motionless  and  incapable  of  producing  impregnation.  Others 
are  active,  but  survive  a  comparatively  short  time.  According  to 
Gary,  the  production  of  the  immature  cells  is  an  effort  on  the  part 
of  the  testes  to  sunplv  an  abnormal  demand,  and  their  presence 
indicates  that  the  fertility  of  the  semen  is  impaired. 

The  deformities,  which  may  be  divided  into  cephalic  and  caudal 
groups,  are  also  the  product  of  defective  spermatogenesis,  or  they 


33 

represent  a  degenerative  process  induced  possibly  by  abnormali- 
ties of  the  fluid  environment.     It  is  rather  difficult,  however,  to 
distinguish  between  deformities  and  immature   types.      The   two 
most  common  cephalic  deformities  are  Avhat  might  be  called  macro 
and  microcephalic  forms.    In  the  former,  the  head  is  enlarged  to  a 
greater  or  less  extent,  it  is  usually  defective  in  staining  qualities, 
and  its  outline  is  indistinct,  due  to  degeneration  of  the  covering 
membrane.     This  type  is  seen  in  Fig.  32.     Also  the  shape  of  the 
head  is  usually  abnormal,  being  either  quite  rounded,  loifg  and 
narrow,  or  short  and  very  broad.     Cells  with  protoplasmic  ap- 
pendages,  though   they   are    more    properly   an   immature  type, 
occasionally  give  the  head  a  greater  volume.    Microcephalic  sperms 
vary  from  those  slightly  smaller  than  normal  to  those  in  which  the 
head  is  represented  by  a  slight  knob.     In  some  cells,  the  head  is 
small  and  round,  in  others,  short  and  stubby,  while  another  type  is 
normal  in  outline  but  diminutive  in  size.     These  forms  likewise 
are,  as  a  rule,  deficient  in  staining  qualities,  and  are  undoubtedly 
degeneration  forms,  occurring  either  as  the  result  of  faulty  de- 
velopment, or  degeneration  subsequent  to  their  formation.     Gary 
believes  they  are  degeneration  types  because  in  the  majority  of  cells 
the  tail  is  apparently  fully  formed,  and  in  the  normal  process  of 
evolution  the   tail   is  the  last  part  of  the   cell  to  be   exhibited. 
Double  headed  forms  are  quite  rare,  but  they  nevertheless  appear 
at  times.    Their  significance  is  difficult  to  explain.    Another  very 
frequent  deformity  of  the  head  is  a  marked  constriction  at  the 
posterior  part  so  that  it  is  the  shape  of  a  pear  or  top  as  in  Fig.  27. 
In  some,  the  head  is  otherwise  normal  in  size.  Avhile  in  others  it  is 
much  elongated,  as  in  Fig.  28,  or  considerably  atrophied.     A  con- 
striction at  the  middle  of  the  head,  as  in  Fig.  29,  is  not  uncommon. 
Both  defects  are  undoubtedlj'-  the  result  of  nuclear  deficiency,  as 
the  nuclenr  part  of  the  head  in  these  cases  is  much  diminished  in 
size,  and  stains  very  deeply  or  not  at  all.    I  have  seen  spermatozoa, 
the  heads  of  which  were  like  an  inverted  cone,  with  a  bulging 
rounded  base.    Other  head.s  are  even  somewhat  contorted  and  bent 
on  themselves,  as  shown  to  some  extent  in  Fig.  26. 

Lender  caudal  deformities,  the  most  frequent  form  encountered 
is  a  thickening  of  the  connecting  piece.  This  may  occur  as  a  uni- 
form thickening,  or  as  a  bulging  appendage.  Rudimentary  de- 
velopment of  the  tail,  the  presence  of  two  poorly  formed  tails,  and 
defective  development  of  the  connecting  piece  occur  rather  infre- 
quently. 

All  these  immature  and  defective  types  are.  as  a  rule,  motion- 
less, and  of  course  incapable  of  producing  fertilization.  Their 
presence  indicates  lowered  fertility  of  the  semen.  Besides  these 
deformities,  there  are  sperms  showing  a  cur\'ature  of  the  tail  at  an 
acute  angle  just  posterior  to  the  neck, —  the  so-called  "  wry  neck." 
Their  significance  is  difficult  to  explain,  but  they  occur  frequently 
in  somen  fixed  and  stained  by  the  same  routine  methods  used  on 
samples  in  which  thev  are  al)sent.  They  probably  are  not  the  re- 
3 


34 

suit  of  the  methods  used  in  fixing  and  staining.  Some  think  they 
are  slightly  immature  types,  or  that  the  condition  is  produced  by 
abnormal  contractions  of  the  tail.  The  majority  of  sperms,  ho^y- 
ever,  especially  those  from  highly  fertile  bulls,  do  not  show  this 
type  at  all. 

The  most  common  changes  in  the  spermatozoa,  are  those  in 
which  there  is  a  separation  of  the  head  from  the  tail,  and  degenera- 
tion of  the  head  as  e^'idenced  by  reaction  to  stains.  The  separation 
of  the  liead  from  the  tail  always  occurs  at  the  neck,  and  often  is 
associated  with  degeneration  or  abnormalities  of  the  head.  The 
separation,  in  the  majority  of  cases,  indicates  some  lowering  of 
vitality  in  the  elements,  altliough  in  many  instances  traumatism 
produced  in  making  smears  or  collecting  the  samples  is  responsible. 
Various  forms  of  abnormal  staining  of  the  head  are  very  common. 
The  cell  membrane,  wliich  is  normally  distinct  and  sharp,  becomes 
blurred  in  outline.  Normally,  the  head  takes  a  good  differential 
stain,  the  anterior  part  staining  lightly,  and  the  posterior  part 
somewhat  deeper.  The  nucleus  is  distinct  in  outline  and  well  de- 
fined. The  lighter  "  inner  body  "  stands  out  in  well  stained  speci- 
mens. As  the  result  of  degeneration,  the  whole  head  may  take 
the  stain  uniformly,  either  slightly  or  much  deeper  than  normal, 
according  to  the  degree  of  degeneration.  The  whole  problem  of 
staining,  however,  depends  very  much  upon  the  methods  used, 
and  the  care  with  which  they  are  applied.  When  a  good  method 
is  obtained,  it  should  lie  adhered  to,  and  used  uniformly  on  all 
specimens.  As  a  rule,  however,  a  certain  amount  of  practice  will 
enable  one  to  differentiate  between  the  sharply  outlined,  clearly 
staining  normal  forms,  and  those  that  show  abnormal  reactions  to 
the  stains. 

Bacteriology 

A  complete  bacteriological  study  was  made  of  the  genital  tracts 
of  fourteen  normal  young  veal  calves  (six  to  twelve  weeks  old), 
four  mature  fertile  IduIIs,  and  sixteen  mature  bulls,  either  sterile  or 
impotent  to  some  degree.  Together  with  these,  the  tracts  of  eleven 
aborted  fetuses,  seven  calves  dying  from  calf  infections  (scours  or 
]:)neumonia).  and  sixteen  bulls  slaughtered  at  an  abattoir  were 
studied  Ijacteriologically.  Occasionally,  studies  were  made  of  indi- 
vidual seminal  vesicles  or  testes,  when  these  parts  alone  were 
brouglit  or  sent  in.  The  history  of  the  abattoir  animals  was,  of 
course,  quite  indefinite  or  entirely  negative.  On  the  killing  floor, 
many  tracts  could  be  studied  for  pathological  changes,  but  in  the 
bacteriological  work  it  was  difficult  to  care  for  more  than  two 
tracts  on  each  visit. 

The  results  of  the  bacteriological  examinations  are  given  in  the 
appended  ta])les,  in  which  the  tracts  are  divided  into  six  groups. 
The  results  in  Oroup  I,  consisting  of  normal  veal  calves,  indicate 
that  the  genital  organs  of  young  male  calves  are,  under  normal 
conditions,  free  from  bacteria.  Carpenter  (9)  obtained  like  re- 
sults in  examining  tlie  genital  tracts  of  heifer  calves..    The  cul- 


35 

tures  made  from  the  seminal  vesicles  and  testes  of  all  these  veal 
calves  were,  with  two  exceptions,  negative.  Both  seminal  vesicles 
of  one  tract  and  one  of  another  yielded  cultures  of  Staphylococcus 
all)  us. 

Adult  bulls  of  known  fertility  were  naturally  difficult  to  obtain, 
only  the  four  animals  in  Group  II  being  available  for  examination. 
Two  of  these  (Nos.  1  and  2)  were  from  the  experimental  herd 
kept  by  the  department,  and  at  all  times  had  a  good  breeding 
history.  The  other  two  were  good  breeders,  but  were  slaughtered 
because  of  poor  pedigrees.  Bull  1,  raised  in  the  department  herd, 
had  a  severe  attack  of  scours  when  a  few  weeks  old,  while  the  calf- 
hood  history  of  the  other  is  not  known,  he  having  been  purchased 
after  reaching  sexual  maturity.  The  cultures  from  the  genital 
organs  of  the  former  (Bull  1)  were  entirely  negative,  exce{)t  those 
from  the  left  epididymis  and  scrotal  sac,  which  yielded  growths  of 
Streptococcus  viridans.  All  tlie  organs  of  the  tract  from  this  ani- 
mal were  normal,  except  foi-  the  fact  that  numerous  strands  of 
connective  tissue  extended  from  the  serous  covering  of  the  tail  of 
both  epididymes  to  the  adjacent  part  of  the  parietal  layer  of  the 
tunica.  The  tract  of  the  other  failed  to  show  any  organisms.  The 
only  evidence  of  any  abnormality  was  the  presence  of  the  same 
connective  tissue  strands  on  the  tail  of  the  epididj-mis,  as  in  the 
first  tract.  The  other  two  Ijulls  gave  negative  cultures  from  all 
parts. 

Of  the  sixteen  bulls  in  Clronp  TIT.  slaughtered  at  abattoii-s,  and 
in  which  no  history  was  available,  eight  failed  to  show  the  pres- 
ence of  any  organisms  in  their  genitalia.  Of  the  others,  the  vesicles 
yielded  cultures  of  Stapliiilococcus  alhus  nine  times,  and  strepto- 
cocci four  times,  Stapltjflococcus  aJhus  was  i-ecovered  once  from 
the  prostate,  and  once  fi'om  Cowper's  glands.  The  testes  gave  cul- 
tures of  staphylococci  in  two  cases,  and  Bact.  ahortum  in  one.  No 
observable  anatomical  changes  accompanied  the  presence  of  the 
Rang  bacillus  in  this  case.  The  epididymes  showed  growths  of 
staphylococci  fi^'e  times,  and  streptococci  on  three  occasions. 
Streptococci  were  isolated   fi"om  the  scrotal  sacs  of  eight  testes. 

The  results  in  Group  IV  (aborted  fetuses)  show  that  bacteria 
are  often  present  in  the  seminal  vesicles  or  testes  of  these  animals. 
As  a  rule,  however,  the  organisms  are  identical  with  those  isolated 
from  the  l)lood  or  other  jiarts  of  tlie  animal.  This  is  to  be  ex- 
pected, however,  for  liecause  of  the  feeble  resistance  of  the  fetus 
to  any  infection,  the  organisms  circulating  in  the  bloo<l  may  be 
isolated,  as  a  rnle.  from  many  ditferent  organs  and  tissues.  All 
samples  of  blood  set  with  Bact.  ahortum  antigen  were  negative, 
irrespective  of  whether  or  not  the  organism  was  recovered  from 
the  blood  or  other  tissue^.  This  is  in  accordance  with  the  findings 
of  Garpenter  in  the  female  fetus. —  the  resistance  is  so  feeble  that 
few  or  no  antibodies  are  formed  to  combat  any  existing  infection. 
Bact.  ahortum  was  recovered  in  two  cases  from  the  vesicles,  and  in 
four  cases  from  the  testes,  but  in  each  instance  the  same  organism 
was  present  in  the  blood  or  otjier  tissues  of  the  body. 


36 

The  results  from  the  tracts  of  the  calves  dying  of  calf  infec- 
tions are  given  in  Group  V,  and  show  tliat  five  were  negative.  The 
other  two  showed  B.  coli,  staphylococci,  and  streptococci,  in  the 
organs  indicated  by  the  chart. 

In  Group  VI,  the  mature  infertile  or  sterile  bulls,  there  was  a 
comparatively  wide  variation  in  the  type  of  organisms  encoun- 
tered, but  streptococci  and  micrococci  were  the  most  common  in- 
vaders. In  the  order  of  the  frequency  of  infection,  the  organs 
would  be  enumerated  as  follows:  Vesicles,  epididymis  (usually 
the  tail),  scrotal  sac,  testes,  prostate,  and  Cowper's  glands.  The 
first  three  parts  mentioned  usually  contained  bacteria.  A  strep- 
tococcus Avas  the  usual  invader  of  the  scrotal  sac,  and  very  prob- 
abl}^  was  the  cause  of  the  connective  tissue  tufts  and  strands  so 
frequently  seen.  The  vesicles  and  epididymes  gave,  in  the  order 
of  the  frequency  of  their  occurrence,  staphylococci,  streptococci, 
B.  coll,  and  Ps.  pyocijaneus.  The  streptococci  were  usually  of 
the  viridans  group,  though  a  few  were  hemolytic,  and  two  strains 
were  indifferent  to  blood.  The  testes  gave  growths  in  only  eleven 
instances, —  staphylococci  eight  times,  streptococci  two  times,  and 
an  unidentified  rod  once.  The  prostate  yielded  staphylococci  twice 
and  Cowper's  gland  once. 

As  emphasized  previously,  the  vesicles  and  tail  of  the  epididymis 
are  most  subject  to  infection  and  degenerative  changes.  At  the 
same  time,  they  are  intimately  connected  with  the  secretion  of 
the  semen.  Once  the  epididymis  becomes  infected,  there  is  nothing 
to  prevent  the  organisms,  together  with  inflammatory  products, 
from  being  mixed  with  the  semen  and  ejaculated  during  coitus. 
Also  in  the  vesicles,  unless  the  inflammation  is  so  severe  as  to 
occlude  the  excretory  duct,  the  organisms  are  mixed  with  the 
vesicular  secretion,  which  is  emptied  into  the  urethra  during 
ejaculation.  Carried  along  v/ith  the  bacteria,  are,  of  course,  toxic 
products,  degenerated  cells,  and  the  otherwise  altered  secretion  of 
the  glands.  One  interesting  case  noted  was  that  of  a  bull  that 
had  passed  from  a  state  of  fertility  to  that  of  complete  sterility 
during  a  period  of  two  months.  The  semen  was  semi-fluid,  green- 
ish yellow  in  color,  and  contained  a  very  few  non-motile  sperma- 
tozoa. Post  mortem  examination  showed  that  the  vesicles  had 
undergone  abscess  formation  and  that  they  contained  yellowish 
green  material  similar  to  that  which  had  been  discharged  during 
copulation.  Streptococcus  hemolytic  us  and  Ps.  jyjfocyaneus  were 
isolated  from  both  vesicles,  and  from  the  semen.  Micrococcus  alius 
was  isolated  in  nearly  all  cases  of  vesiculitis  and  was  often  asso- 
ciated with  Streptococcus  viridaus  or  hemolyticus. 

Bacteriological  studies  of  the  semen  are,  on  the  Avhole,  more  or 
less  unsatisfactory,  due  to  the  present  difficulty  in  obtaining  sam- 
ples free  from  any  chance  of  contamination.  In  most  of  the  ab- 
normal bulls,  bacteria  of  various  types  were  isloated  from  the 
semen,  most  of  which  agreed  culturally  with  those  later  isolated 
from  the  internal  genital  organs  of  the  same  tracts.  The  method 
of  culturing  consisted  of  douching  the  prepuce  of  the  bull  and 


37 

vagina  of  the  female  with  sterile  saline  solution  before  breeding. 
Samples  of  vaginal  mucus  were  taken  before  service,  and  the  flora 
compared  to  that  after  douching.  This  method  of  douching  pro- 
duced vaginal  samples  relatively  free  from  bacteria,  at  least  so 
much  so  that  the  post  coital  fluid  demonstrated  that  many  organ- 
isms must  have  been  introduced  from  without.  AA^hether  or  not 
they  came  in  with  the  semen  is  problematical,  but  in  all  probability 
this  was  the  method  of  introduction. 

I  have  so  far  failed  to  obtain  Bad.  ahortum  from  the  tract  of  an 
adult  animal,  either  by  direct  culture  or  guinea  pig  inoculation, 
except  from  the  testicle  of  one  abattoir  bull.  The  agglutination 
tests  with  Bad.  ahortum  antigen  were  all  negative,  except  for  tAvo 
abattoir  bulls.  The  results  so  far  obtained  would  seem  to  indicate 
that,  in  accordance  with  the  findings  of  other  workers,  the  Bang 
organism  seldom  invades  the  male  genital  tract,  or  does  not  thrive 
there  after  its  introduction.  Schroeder  (12)  and  others,  have, 
however,  on  various  occasions,  recovered  the  organism  from  the 
bull,  and  the  former  author  even  states  that  it  invades  the  vesicles 
and  is  eliminated  with  the  semen. 

Discussion 

A  complete  discussion  of  those  factors  which  have  a  bearing  on 
reproduction  and  fertility  in  an  animal,  includes  not  only  a 
thorough  study  of  the  genital  tract,  but  an  appreciative  considera- 
tion of  various  extrinsic  factors.  The  effect  of  environment  has 
long  been  known  to  have  a  marked  influence  upon  breeding,  par- 
ticularly Avith  reference  to  animals  in  domestication.  Diet,  though 
long  relegated  to  a  minor  phase  of  the  question,  has,  Avithin  recent 
years,  come  to  be  a  matter  of  prime  importance  AA'ith  regard  to  its 
bearing  upon  the  entire  body  metabolism.  The  endocrine  organs 
preside  over  and  regulate  the  gi'owth  and  functioning  of  the  geni- 
tal organs  from  the  earliest  embryonic  stage  to  the  cessation  of 
sexual  life.  Any  derangement  in  one  results  in  functional  or 
organic  changes  in  the  other.  In  a  given  mating,  aa^c  must  take 
into  consideration  such  factors  as  impediments  to  coitus,  as  well 
as  those  numerous  agencies  in  the  female  AA^hieh  may  interfere  with 
the  union  of  sperm  and  OAaim,  or  AA'ith  the  successful  implantation 
of  the  fertilized  eg^^  in  the  uterus,  and  its  groAvth  and  deA^elop- 
ment  there  till  normal  parturition  takes  place.  Successful  repro- 
duction depends  upon  the  mating  of  sexually  sound  females  to 
equally  sound  males.  Considering  the  A^arious  factors  AA-hich  gov- 
ern reproduction,  sexual  soundness  must  necessarily  depend,  to 
a  large  extent,  upon  a  good  general  condition  of  the  entire  body. 

ExA'iRONMENT:  The  effeet  of  environment  on  fertility  in  the 
bull  is  no  doubt  a  minor  factor.  Cases  in  Avhich  changes  in  en- 
vironment affect  fertility  probal)ly  occur,  liOAvcA-er,  particularly 
when  fear  and  other  psychic  disturbances  play  a  part.  Marshill 
(29)  states:  "Tt  Avould  seem  probable  that  failure  to  breed 
among  animals  in  a  strange  enA'ironment  is  due  not,  as  has  been 


38 

sxinrtrested,  to  any  toxic  influence  on  the  organs  of  generation,  but 
to  the  same  causes  as  tho.se  which  restrict  breeding  in  a  state  of 
nature  to  certain  particular  seasons,  and  that  the  sexual  instinct 
can  only  be  called  into  play  in  response  to  certain  stimuli, —  the 
existence  of  which  depends  to  a  large  extent  upon  appropriate 
seasonal  and  climatic  changes." 

Diet:  Under  tliis  heading  we  may  include  not  only  the  effect 
of  deficient  food,  but  also  constitutional  disorders,  as  a  result  of 
"which  the  oi'gans  of  generation  and  those  glands  guarding  their 
function  I'cceive  insufticient  nourishment.  It  is  a  well  known  fact, 
and  long  has  l>cen.  that  animals  fail  to  breed  when  they  are  in  a 
run  down  condition  or  when  they  are  fed  a  deficient  diet.  Gary 
(35),  quoting  Hagner,  states  that  the  virility  of  the  spermatozoa 
is  often  in  direct  proportion  to  the  general  physical  condition  of 
the  patient. 

Reynolds  (di)  emphasizes  the  fact  that  it  is  an  established 
l^rinciple  among  animal  breeders  that  a  high  protein  diet  in  both 
sexes  is  essential  to  full  fertility.  "  Oligospermia  with  defieient 
vitality  of  the  spermatozoa  is  not  infrequently  found  from  con- 
stitutional disorders.  It  can  easily  be  demonstrated  in  animals 
that  both  low  diet  and  conditions  of  life  that  produce  a  nervous 
excitable  state  are  attended  by  oligospermia."  Animals  that  are 
closely  confined,  those  that  are  over-fat  (show  animals),  as  well 
as  those  fed  a  deficient  ration  very  frequently  fail  to  breed,' but 
exercise  and  change  of  diet  soon  overcome  the  impotency. 

Dutscher,  Hart,  Steenbock.  and  other  biological  chemists  have 
done  extensive  work  to  show  the  essential  importance  of  vitamines 
and  minerals  in  the  diet.  Their  results  indicate  that  animals  can- 
not thrive  and  breed  normally  when  fed  a  diet  composed  solely  of 
the  products  of  one  plant.  There  must  be  variety,  and  there  must 
be  not  only  a  correct  nutritive  ratio,  but  the  mineral  and  vitamine 
content  must  be  present  as  well.  Cows  fed  on  the  products  of  one 
plant  often  failed  to  ])reed,  and  if  conception  occurred,  it  in- 
variably^ resulted  in  a  premature  birth,  or  the  birth  of  weak  and 
poorly  nourished  calves.  The  work  of  these  authors  is  funda- 
mental, and  Ijrings  out  many  important  points.  Is  it  not  probable 
that  the  deficient  diet  results  in  Aveakened  tissues  which  are  easier 
proy  to  the  invasion  of  bacteria? 

Macomber  and  Reynolds  (39)  experimented  upon  white  rats 
to  determine  the  effect  of  defective  diet  as  a  cause  of  .sterility. 
They  call  attention  to  the  confusion  caused  by  the  application  of 
the  term  sterility  to  most,  or  all,  infertile  matings.  They  believe 
that  failure  of  reproduction  is,  in  fact,  the  result  of  cleereased 
fertility  rather  than  of  actual  sterility  on  the  part  of  the  two  indi- 
viduals concerned.  "  There  are  certainly  a  large  number  of  in- 
fertile matings  which  are  purely  functional  and  due  to  physiologic 
alterations  or  local  conditions.  Such  physiological  alterations  more- 
over coexist  in  the  sterilities  of  ])atholoo:ic  origin  and  when  un- 
recognized and  consequently  unremedied,  undoubtedly  explain  a 


39 

large  proportion  of  tlu'  continued  infertilities  after  operation." 
Is  it  possible  for  a  bull  to  be  infertile  to  the  cows  in  his  herd 
that  have  been  fed  a  deficient  diet,  and  at  the  same  time  to  breed 
well  when  mated  to  animals  outside  this  herd  ?  This  is  rather  im- 
probable in  practice,  but  there  is  always  the  possibility  of  its 
oceurrenee.  In  tlie  experimental  work,  white  rats  were  used:  one 
strain  from  tlie  \Vistar  Institute  with  a  fertility  of  about  65  per 
cent,  and  the  other  from  a  Dr.  Castle's  strain  with  a  fertility  of 
about  90  per  cent.  The  "Wistar  rats  were  fed  in  groups,  each 
group  receiving  a  diet  deficient  in  a  certain  sul)stance :  calcium, 
protein,  or  fat  soluble  vitamine.  To  this  grou])  was  added  a  diet 
deficient  in  both  calcium  and  protein  (war  diet).  These  diets 
reduced  the  fertility  of  the  groups  from  the  original  65  per  cent, 
to  55,  ?)1.  and  14  per  cent  respectively.  It  delayed  the  appearance 
of  fertilitv  in  voung  rats  raised  on  these  diets,  and  lowered  its 
degree  in  the  mature  animals.  Most  of  these  rats,  however,  though 
infertile  to  each  other,  bred  promptly  Avhen  mated  to  the  Castle 
rats  of  known  fertility.  This  demonstrates  clearly  that  relative 
infertility  of  given  matings  does  occur.  One  interesting  feature 
of  the  work  is  the  fact  that  in  the  matings  on  the  single  deficiency 
diets,  four  deliveries  of  macerated  fetuses  occurred  and  there  were 
two  more  in  eight  deliveries  from  those  reared  on  the  war  diet. 
Xo  cases  of  this  kind  had  previously  occurred  in  this  strain,  which 
had  1)ecn  under  observation  for  several  years.  Does  this  throw 
any  light  upon  the  cause  of  macerated  fetuses  in  cattle?  Micro- 
sco]iically  the  testes  and  ovaries  of  these  infertile  rats  showed  no 
observable  changes,  a  fact  which  is  of  great  importance  to  bear  in 
mind. 

Williams,  in  his  book  on  disease  of  the  genital  organs,  brings  out 
quite  clearly  the  relation  of  defective  diet,  overfeeding,  and  lack 
of  exercise,  to  reproductive  efficiency. 

Xovarro  (40)  observed  that  pigeons  fed  on  a  diet  without  vita- 
mine  B  showed  degeneration  of  the  seminal  epithelium,  with 
hypertrophy  and  hyperplasia  of  the  interstitial  cells  of  the  testis. 
Another  author  (Allen)  .showed  that  reduction  in  the  quantity  of 
water-soluble  vitamine  in  the  diet  of  rats  resulted  in  total  degenera- 
tion of  all  the  germ  cells,  but  it  did  not  interfere  witli  gi'owth  and 
development  in  other  respects. 

The  observations  of  Williams  (41),  in  a  pure  bred  beef  herd  in 
Hawaii,  clearly  demonstrate  the  intimate  correlation  between  poor 
fodder  as  the  result  of  extreme  drought,  and  the  accentuation  of. 
or  increased  susceptibility  to,  genital  infections,  as  demonstrated 
by  clinical  findings.  The  genital  disorders  started  soon  after  the 
onset  of  the  drought,  and  immediately  took  a  downward  trend 
with  the  advent  of  the  rainy  season. 

Judging  by  the  work  quoted,  we  will  observe  tliat  deficient  diet, 
though  it  does  not  always  affect  the  general  health,  has  a  ]irofnund 
effect  upon  the  genital  organs  of  l)oth  sexes,  associated  with  dis- 
turbances of  spermatogenesis  in  the  male.    In  most  debilitated  and 


40 

weakened  conditions  of  the  male,  spermatogenesis  ceases  or  is 
markedly  defective.  We  must,  undoubtedly,  explain  this  fact 
upon  the  ground  of  deficient  nouHshment  to  the  reproductive 
organ.s  or  possibly  the  endocrines.  The  vitamines  have  been 
termed  nuclear  nourishers,  and  their  absence  probably  results  in 
nuclear  deficienc}'. 

ExDOCRiNES:  Bell  (42)  emphasizes  the  fact  that  not  only  the 
structure  but  also  the  function  of  every  part  of  the  body  is  in 
close  correlation  with  the  rest.  "  This  is  essentially  true  of  the 
ductless  glands:  the  shadow  of  their  influence  is  over  all." 
Further  he  states  that  when  we  remember  that  the  individual 
exists  to  perpetuate  the  species,  it  is  not  difficult  to  realize  that 
the  metabolic  factors  concerned  in  reproduction  are  the  same  as 
those  related  to  the  individual  metabolism.  It  follows,  therefore, 
that  the  ductless  glands  which  regulate  the  individual  metabolism 
concern  equally  the  reproductive.  Brown  (43),  discussing  the 
same  subject  brings  out  the  generalization  that  the  sympathetic, 
since  it  is  the  most  primitive  part  of  the  nervous  system,  is  closely 
associated  with  the  endocrine  system,  a  still  more  elemental  means 
of  communication  in  the  body.  Also  since  specialized  reproductive 
cells  appear  before  the  nervous  system,  the  organs  of  reproduction 
remain  closely  associated  with  the  older  chemical  reactions  now 
specialized  in  the  endocrine  glands.  "  The  endocrine  glands,  the 
reproductive  organs,  and  the  sympathetic  nervous  system,  there- 
fore, remain  as  a  basic  tripod,  and  it  is  not  likely  that  a  disturb- 
ance will  occur  for  long  in  one  limb  without  affecting  the  other 
two."  The  former  author  believes  that  the  gonad  keeps  the  other 
ductless  glands  informed  of  the  needs  of  the  genital  tract,  they  in 
turn  influencing  the  general  metabolism.  Jump  (44)  states: 
"  We  are  therefore  justified  in  believing  that  there  is  a  correlation 
of  function  between  these  (endocrine)  glands  and  that  some  cases 
of  sterility  are  probably  due  to  a  derangement  of  this  correlation." 
Biedl  (45 ")  concludes:  "  There  appears  to  be  an  intimate  anatomi- 
cal and  physiological  interrelationship  between  the  different  blood 
glands  which  is  manifested  clinically  by  the  fact  that  the  patho- 
logical disturl)ance  of  one  gland  is  accompanied  by  symptoms  point- 
ins:  to  the  functional  derangement  of  one  or  more  of  the  others. 
Knowing,  as  we  do,  the  many  sided  interactivity  which  subsists  be- 
tween the  different  internal  secretory  organs,  itTs  readily  con- 
ceivable that  isolated  diseases  of  single  organs  of  this  group  are 
very  much  rarer  than,  at  the  first  glance,  they  would  appear  to  be. 
Tn  the  present  state  of  our  knowledge,  the  only  course  of  investiga- 
tion which  is  open  to  us  is  to  start  with  the  known  results  of  the 
functional  derangement  of  any  organ,  and,  by  following  these  up, 
to  seek  the  primary  link  in  the  pathological  chain." 

Most  workers  seom  to  agree  that  a  special  connection  exists  be- 
tween the  normal  function  of  the  adrenal  cortex  and  the  sexual 
organs.     Tumors  of  the  former  are  usually  associated  with  sex 


41 

abnormalities,  and  feeding  young  animals  the  gland  substance 
seems  to  stimulate  growth  or  the  testes. 

Many  arguments  have  been  brought  forward  to  show  that  the 
prostate  produces  an  internal  secretion.  It  is  a  well  known  fact 
that  this  organ  atrophies  after  castration,  and  enlarges  as  the  sex 
life  dwindles.  As  has  been  previously  stated,  Serrlach  and  Pares 
reached  the  conclusion  that  the  gland  prodvices  an  internal  secre- 
tion which  controls  the  testicular  functions  and  regulates  the  pro- 
cess of  ejaculation.  Also  they  state  that  if  the  prostate  is  re- 
moved, spermatozoa  are  no  longer  produced  in  the  testes,  and  that 
the  secretory  activity  of  the  accessory  genital  gland  ceases.  The 
secretion  is,  at  any  rate,  a  stimulus  to  the  internal  secretion  of  the 
testis. 

The  thyroid  bears  a  distinct  biological  relationship  to  the  sexual 
glands.  Removal  of  the  gland  results  in  imperfect  development  of 
the  gondas.  infantilism,  and  general  torpor.  Bell  (42)  believes 
that  the  association  between  ^:he  thyroid  gland  and  the  genitalia  is 
as  intimate  as  the  relation  of  the  pituitary  to  the  genital  functions. 

Of  all  the  endocrines,  perhaps  the  anterior  lobe  of  the  hypophysis 
is  in  most  intimate  correlation  with  reproduction.  Castration 
results  in  hypertrophy  of  this  organ,  while  removal  of  the  anterior 
lobe  usually  leads  to  death.  In  those  cases  in  which  death  does 
not  ensue,  it  results  in  genital  atrophy,  stunting,  and-  reduc- 
tion of  sexual  activity.  In  young  animals,  spermatogenesis  ceases 
entirely  even  after  partial  extirpation  of  the  anterior  lobe,  Biedl 
(45)  states  that  "  in  disease  of  the  hypophysis,  derangement  of 
sexual  acti^-ity  occurs  very  early  in  the  course  of  the  disease.  sho'v\Ti 
in  women  by  the  cessation  of  menstruation,  and  in  men  by  im- 
potence," 

The  thymus,  as  is  well  known,  is  quite  intimately  associated 
with  the  development  of  the  genital  organs.  Its  normal  disap- 
pearance is  always  associated  with  the  development  of  sexual 
maturity  in  the  individual.  Hewer  (46)  conducted  experiments  to 
ascertain  the  effect  of  thymus  feeding  on  the  activity  of  the  re- 
productive organs  in  the  rat.  She  concludes  in  part :  ' '  Male  rats 
appear  more  susceptible  to  the  influence  of  thymus  feeding  than 
female  rats.  With  moderate  doses  of  thymus,  sexual  maturity  in 
the  animals  treated  is  delayed,  a  phenomenon  which  is  attributed 
to  delayed  development  of  the  testis.  AYith  large  doses  of  th\nnus, 
in  the  male,  the  testis  is  structurally  affected :  in  the  young  animal 
in  the  direction  of  retardation  of  development,  in  the  mature  ani- 
mal in  the  direction  of  degeneration.  This  degeneration  is  con- 
fined to  the  testes.  In  the  degenerating  testis,  cells  of  Sertoli  ap- 
pear to  be  absent :  the  spermatogonia  are  present,  also  dividing,  and 
may  lie  free  in  the  lumen  of  the  tubule ;  spermatids,  many  with 
abnormal  nuclei,  are  shed  into  the  lumen  in  large  numbers ;  sper- 
matozoa are  practically  nbsent.  In  the  later  stages,  only  a  few 
dividing  spermatogonia  appear  among  the  debris  of  the  other  un- 
recognizable cells  of  the  tubide.    In  the  epididymis  Avhich  itself  is 


42 

normal,  when  the  testis  is  showing  degeneration,  ver^"  few  spei*ma- 
tozoa  appear,  in  the  later  stages  none.  Many  spermatids  are  pres- 
ent in  various  stages,  and  some  spermatocytes.  Animals  in  the 
hyper-thymic  condition  appear  to  be  sterile." 

The  foregoing  references  will,  I  hope,  serve  to  bring  out  the  facts 
that  environment  and  diet,  together  Avith  the  general  body  meta- 
bolism and  the  endocrines,  have  a  more  or  less  profound  effect 
upon  the  development,  growth,  and  functioning  of  the  genital 
system.  In  the  experiments  it  has  been  shown  that  sterility  is 
not  necessarily  accompanied  by  any  apparent  microscopical 
changes  in  the  gonads,  or  even  at  times  in  the  general  body  health. 
Nor  can  w^e  exclude  impotency  of  the  male  entirely  even  when  the 
spermatozoa  are  normal  in  shape,  and  motility.  Carnett,  and 
others  (38),  years  ago  stated:  "  Indeed,  there  is  abundant  clini- 
cal proof  to  the  effect  that  systemic  conditions  which  have  no  ap- 
preciable effect  upon  the  motility  or  conformation  of  the  sperma- 
tozoa materially  interfere  with  reproductive  power."  The  entire 
complex  genital  system  is  inseparably  linked  up  with  the  body  as  a 
whole,  a  fact  which  we  must  bear  in  mind  at  all  times. 

Impediments  to  coitus  may  be  due  to  great  difference  in  the  size 
of  the  two  mated  individuals,  psychic  disturbances,  or  inability 
to  protrude  the  penis.  Williams  (17)  mentions  several  physical 
impediments,  as  deformity  of  the  limbs  or  feet,  sore  feet,  overload- 
ing of  the  rumen,  obesity,  fear  of  falling,  and  paralysis.  Coitus 
may  be  somewhat  delayed,  or  even  not  performed  as  the  result  of 
a  severe  inflammation  with  sensitiveness  of  the  penis  or  prepuce. 
Occasionally  tumors  of  the  penis  are  encountered  which  may  inter- 
fere with  prolusion  of  the  penis,  or  its  entrance  into  the  vagina. 
Not  infrequently  the  penis  is  rendered  incapable  of  protrusion  as 
the  result  of  inflammatory  adhesions,  tuberculosis  of  the  preputial 
lymph  glands,  etc. 

Excessive  sexual  use,  within  certain  limits,  probably  has  not,  in 
itself,  any  material  permanent  effect  upon  the  reproductive  capac- 
it}-.  Tlie  frequency  with  which  bulls  used  to  excess  break  down 
sexually,  is  probably  due  to  the  devitalizing  effect  upon  the  tissues 
of  tlie  genital  organs,  this  opening  the  way  to  bacterial  invasion 
and  other  destructive  influences.  Over-use  is  probably  not  danger- 
ous, unless  continued  over  long  ]ieriods.  but  at  the  same  time  it 
offers  greater  opportunity  for  infection  to  be  introduced  into  the 
body  from  intercourse  with  large  numbers  of  females.  Lloyd- 
Jones  and  Hays  (47)  carried  on  very  interesting  experiments  on 
the  influence  of  excessive  sexual  activity  of  male  rabbits  on  the 
properties  of  the  semen.  Their  plan  was  to  mate  male  rabbits  in 
quick  succession,  and  study  the  character  of  the  semen  on  the 
first  service,  and  every  fifth  service  thereafter.  The  safe  limit 
was  twenty  services  in  three  hours.  As  would  be  expected,  the 
volume  of  the  semen,  after  the  first  few  services,  became  gradually 
reduced  in  amount.  "  In  rapidly  successive  services,  the  semen 
becomes  less  viscous  and  tends  to  lose  its  charactertistic  milky 


43 

appearance  until  at  the  twentieth  service,  when  the  fluid  is  thin 
and  watery,"  It  seemed  as  though  there  was  a  well  marked  reduc- 
tion in  the  number  of  spermatozoa  per  cubic  centimeter  in  the  ad- 
vanced servdces.  Successive  copulations  also  resulted  in  a  marked 
decrease  in  the  number  of  motile  spermatozoa,  together  with  a 
sliorter  duration  of  perceptible  vitality.  The  certainty  of  produc- 
ing impregnation  at  the  same  time  became  less  and  less.  "This 
reduction  in  the  per  cent  of  effective  matings  when  the  male  is 
sexually  overworked  is  recognized  hy  those  engaged  in  animal 
breeding  as  one  of  the  most  noticeable  and  universal  concomitants 
of  heavy  sexual  service." 

In  another  paper,  these  same  authors  studied  the  effect  of  sexual 
excess  upon  the  character  of  the  offspring.  In  part,  they  conclude : 
''Ba'  no  means  thus  far  used  has  any  inferiority  of  i)rogeny  from 
the  heavy  sexual  service  been  discovered.  They  are  fully  equal  if 
not  superior  to  progeny  from  very  light  service  of  male." 

Infection  is  without  doubt  the  greatest  single  factor  capable  of 
producing  functional  and  anatomic  changes  resulting  in  varying 
degrees  of  imi)oteiicy  and  sterility.  The  changes  produced  range 
from  the  addition  of  the  toxic  products  of  bacterial  growth  to  the 
seminal  fluid,  to  the  complete  destruction  of  the  parenchymatous 
tissue  of  one  or  more  of  the  contributing  sextial  glands.  Anatomic 
changes  are  l)y  no  means  essential  to  the  ])roduction  of  lowered 
fertility.  As  has  been  previously  stated,  the  work  on  veal  calves 
indicates  that  the  genital  organs  of  young  bulls  are  normally  free 
from  bacteria.  Likewise  in  normal  adult  animals,  the  bacterial 
content  of  the  genital  organs  is  as  a  rule  low  or  negative.  It  is 
possil)le  that  a  certain  flora  is  normal  for  the  tract  at  sexual 
maturity,  as  in  several  other  organs  of  the  body,  but  under  the 
strain  of  sexual  excess,  defective  diet,  or  other  weakening  in- 
fluences, the5-e  organisms  may  become  pathogenic.  Streptococci  and 
staphylococci  have  at  times  been  found  in  apparently  normal  parts 
of  the  body,  and  at  other  times  they  are  found  associated  with 
severe  pathological  lesions  in  the  genital  tract.  The  degree  of 
pathogenicity  is  of  course  difficult  to  determine,  except  as  we  find 
the  bacteria  associated  with  abnormal  conditions.  Carpenter  (9), 
however,  injected  streptococci  into  the  genital  tracts  of  female 
calves  and  produced  lesions  resem])ling  very  closely  those  from 
which  the  organisms  had  been  isolated  in  adult  sterile  animals.  Per- 
sonally. I  am  inclined  to  believe  that  the  genital  organs  normally 
are  free  from  bacteria,  or  if  any  are  there  they  are  better  able 
to  multiply  under  the  strain  of  devitalization  of  the  tis.sues. 
Bacterial  invasion,  liOAvever,  does  take  ]ilace  fpiito  fi-equently.  l)ut 
the  paths  of  entrance  of  the  organisms  are  .somewhat  proble- 
matical. Hematogenous  origin  is  always  possible,  though  it  is 
rather  difficult  to  definitely  implicate  this  mode  of  entrance.  The 
urethra  is  perhaps  the  easiest  and  most  common  ]iath  for  the 
entrance  of  bacteria,  though  even  here  it  is  not  ])Ossible  to  make 
definite  assertions.     Contiguous  spread  of  infection  from  neigh- 


44 

boring  structures  is  very  probable  iu  some  cases,  particularly  in 
pelvic  peritonitis.  The  Ijaeteriological  results  hardly  bear  out  the 
theory  of  ^Yilliams  that  the  organisms  lie  dormant  in  the  genitalia 
of  the  animals  until  the  advent  of  sexual  maturitj^,  at  which  time 
they  acquire  pathogenic  powers.  On  the  other  hand,  his  clinical 
observations  seem  to  indicate  that  this  may  be  possible.  Calves 
suffering  from  "calf  infections"  frequently  do  harbor  organisms 
in  their  genital  organs,  but  whether  or  not  they  persist  there  till 
sexual  maturity  is  a  matter  of  conjecture.  The  most  logical  theory 
seems  to  be  that  animals  from  herds  in  which  genital  infections 
are  very  severe,  or  those  that  have  had  severe  attacl<s  of  scours 
or  pneumonia,  are  more  susceptible  to  those  infections,  due  to  the 
early  lowering  of  their  vitality.  One  bull  in  the  department  herd 
certainly  had  a  severe  ordeal  as  a  calf,  but  as  a  mature  bull  he 
was  highly  fertile.  Moderate  sexual  use  and  proper  sexual  hygiene 
proljably  had  much  to  do  with  this.  In  the  bull,  infectjon  of 
some  part  of  the  genitals,  during  some  period  of  life,  is  very  con- 
stant, however,  whether  or  not  it  is  productive  of  observable 
changes  in  his  breeding  efficiency.  The  finding  of  the  fine  connec- 
tive tissue  strands  and  tufts  on  the  serous  surface  of  the  tail  of 
the  epididymis  of  practically  all  bulls  examined,  both  sterile  and 
fertile,  indicates  past  or  present  infection  of  the  scrotal  sac.  The 
vesicles  and  tail  of  the  epididymis  are,  as  stated  previously,  the 
most  commonly  invaded  tissues  of  the  tract.  The  testes  are  less 
frequently  involved. 

While  it  is  difficult  to  obtain  irreproachable  proof  that  the  bull 
is  a  disseminator  of  genital  infections,  the  findings  of  clinicians 
quite  clearly  indicate  that  this  is  true,  and  laboratory  methods 
tend  to  support  this  assumption.  Williams  believes  that  not  only 
may  the  bull  infect  the  female  with  organisms  whicli  interfere 
with  the  given  conception,  but  that  he  often  implants  there  organ- 
isms which  interfere  with  future  pregnancies,  and  even  with  the 
life  of  the  individual  in  some  cases.  The  high  abortion  and  ster- 
ility rate  following  the  use  of  certain  sires,  and  the  appearance  of 
characteristic  infections  after  seiwice  to  certain  bulls,  clearlj^  in- 
dicate that  in  all  probability  the  bull  does  eliminate  with  his 
semen  those  organisms  whicli  produce  lesions  in  his  genital  organs, 
and  are  capable  of  infecting  the  female.  W.  L.  Williams  (48) 
cites  the  case  of  a  pure  bred  herd  in  which  breeding  had  pro- 
gressed satisfactorily  until  heifers  had  grown  to  breeding  age  and 
a  second  bull  was  obtained.  "  Some  cows  of  the  old  herd  were 
also  assigned  to  the  young  bull  which  had  not  previously  been  in 
service.  The  cows  bred  to  the  old  herd  bull  continued  to  breed 
normally.  The  cows  and  heifers  bred  to  the  new  bull  conceived 
with  difficulty  or  not  at  all.  Those  which  conceived  mostly 
aborted,  and  those  which  calved  had  metritis  and  retained  fetal 
membranes.  The  two  first  cows  in  which  pregnancy  terminated 
died  of  metritis."  I  have  frequently  had  semen  samples  sent  in 
from  bulls  that  were  not  only  failing  to  get  cows  with  calf,  but 


45 

following-  each  service  the  females  showed  a  severe  vaginitis. 
\V.  W.  ^\  illianis  worked  in  a  herd  in  which  service  to  certain  bulls 
was  in  each  case  followed  by  a  severe  vaginitis  and  cervicitis,  only 
to  be  followed  later  by  a  characteristic  salpingitis. 

Vaginal  smears  taken  before  and  after  service,  in  many  in- 
stances, show  that  in  all  probability  bacteria,  especially  the  strep- 
tococci, were  deposited  there  with  the  semen.  These  results  have 
been  obtained  upon  several  occasions,  at  which  time  the  vagina 
was  usually  douched  prior  to  each  service  with  sterile  saline  solu- 
tion. Streptococci  and  other  organisms  have  been  isolated  from 
the  vaginal  samples  obtained  by  this  method.  In  most  cases,  they 
were  absent  from  samples  taken  before  service.  Extraneous  con- 
tamination, and  error,  must  l^e  taken  into  consideration,  but  the 
results  tend  to  bear  out  clinical  observations  that  the  bull  is  prob- 
ably a  disseminator  of  some  infections  associated  with  the  genital 
organs  of  both  sexes.  At  any  rate,  organisms  have  been  isolated 
repeatedly  from  the  genital  organs  of  the  bull,  of  the  same  bio- 
logical character  as  those  which  are  associated  with  sterility,  abor- 
tion, and  allied  phenomena  in  the  female.  In  the  absence  of 
obstruction  in  any  part  of  the  tract,  there  is  nothing  to  hinder 
infection  from  gaining  access  to  the  seminal  fluid,  and  being  ex- 
creted during  ejaculation. 

Hopper  (18)  states:  "A  diseased  bull  may  manifest  non-fertility 
or  decreased  potency  in  different  ways — by  repeated  service  to 
apparently  normal  females  without  conception,  by  a  high  abortion 
rate  in  females  that  have  been  apparently  normal,  by  characteristic 
infections  following  the  use  of  any  particular  sire,  or  by  abnor- 
malities in  the  breeding  tract  noted  by  rectal  or  physical  pal- 
pation." 

Admittedly,  Bad.  abortum  has  little  affinity  for  the  genitalia 
of  the  bull,  though  Sehroeder  states  that  the  bull  harbors  the 
organisms  in  his  seminal  vesicles  and  that  they  are  eliminated 
with  the  semen.  Other  authors  have  occasionally  isolated  the  or- 
ganism from  the  vesicles,  testes,  or  both.  Sehroeder 's  theory  that 
infection  of  the  female  occurs  indirectly  by  contamination  of  the 
fo<lder  with  the  semen  is  prol)ably  rare  in  occurrence.  The  very 
limited  number  of  cases  in  which  investigators  have  demonstrated 
the  presence  of  the  organism  in  the  male  genital  organs,  and  the 
apparent  immunity  of  the  bull  to  the  bacterium  as  determined  by 
the  agglutination  reaction,  seem  to  indicate  that  he  plays  a  small 
part  in  the  spread  of  this  type  of  infection  in  the  here!.  On  the 
other  hand,  it  would  seem  that  he  is  at  times  intimately  associated 
with  the  spread  of  certain  other  organisms  that  interfere  seriously 
with  herd  reproductivity. 

The  diagnosis  of  infertility  and  sterility  rests  upon  a  thorough 
physical  examination  of  the  genital  organs,  together  with  a  de- 
tailed study  of  the  semen.  The  histoiy  of  the  animal  and  herd 
involved  must  also  be  verj'-  carefully  inquired  into,  especially  the 
part  covering  the  result  of  everj'  serv'ice  by  the  sire  in  question. 


46 

Besides  this,  we  must  always  consider  all  factors  which  have  a 
bearing  upon  the  subject,  remembering  the  phj'siology  of  each 
part,  and  the  role  it  plays  in  reproduction.  Bacteria  gain  entrance 
to  many  parts  of  the  tract,  where  they  multiply  and  probably  add 
toxic  products  to  the  seminal  fluid,  altering  its  biological  char- 
acter and  resulting  in  partial  or  total  destruction  of  the  secretory 
tissues.  The  testes,  epididymes,  vesicles,  prostate,  and  other  parts, 
each  contribute  their  essential  part  to  the  semen,  abnormalities  of 
any  one  of  which,  as  a  rule,  result  in  interferences  with  reproduc- 
tion. If  the  vesicles  are  involved,  we  must  bear  in  mind  just  what 
is  the  part  played  by  their  secretion,  and  what  is  the  probable 
result  if  their  essential  elements  are  not  added  to  the  semen.  In 
like  manner,  we  must  consider  the  prostate,  whose  secretion  stimu- 
lates the  vitality  of  the  sperms,  and  adds  fluid  bulk  to  the  semen. 
Extirpation  of  the  vesicles  or  prostate  alone  results  in  lowered 
fertility,  without  altering  the  sexual  desire,  while  removing  lioth 
glands  produces  total  sterility.  Partial  or  total  destruction  of  the 
parenchyma  of  either  gland  produces  the  same  effect  as  extirpa- 
tion, in  that  its  function  is  altered  or  entirely  absent. 

The  semen  should  be  examined,  not  only  for  the  number  of 
spermatozoa  and  the  percentage  of  those  that  are  motile,  but  for 
the  duration  and  type  of  motion.  Normal  semen,  when  first  ex- 
amined under  the  microscope,  shows  a  field  closely  packed  with 
highly  motile  spermatozoa.  In  every  study  of  the  semen,  however, 
we  must  1)ear  in  mind  the  temperature  and  other  conditions  under 
which  it  has  been  kept  since  emission.  On  the  other  hand,  semen 
from  bulls  of  lowered  fertility  shows  changes  ranging  from  mild 
disturbances  such  as  sluggish  motility  and  a  slight  decrease  in  the 
number  of  sperms  present,  to  aspermia,  or  total  lack  of  motion. 
Normal  semen,  Avhen  compared  with  abnormal  specimens,  as  a  rule 
presents  distinct  differences,  either  in  motility,  staining  properties, 
or  structure  of  the  spermatozoa.  Impotent  bulls,  however,  may 
show  at  times  few  or  no  ol)servable  changes  in  their  genital  organs. 
The  only  assumption  hei-e  is  that  the  condition  probably  is  of 
endocrine  origin,  or  is  some  functional  disturbance.  Of  oligo-, 
spermia  Reynolds  states:  ''Oligospermia,  Avith  normal  motility 
and  vitality,  is  not  al)so]ute  sterility,  l)ut  is  of  high  importance 
because  the  percentage  of  destruction  of  spermatozoa  during  their 
passage  through  the  genital  canal  of  the  female  is  so  enormous  that 
the  possibility  of  impregnation  by  semen  which  starts  out  with  a 
deficient  number  is  always  poor.  When  the  genitals  of  the  female 
partner  are  in  a  condition  which  is  even  moderately  hostile  to  the 
spermatozoa,  impregnation  by  such  semen  becomes  so  unlikely  as 
to  be  not  even  a  probability." 

Motility  may  be  lacking  in  a  small  number,  its  absence  may  be 
observed  in  a  large  percentage,  or  even  in  all  those  in  the  field. 
as  in  necrospermia.  On  the  other  hand,  the  motility  may  be 
sluggish  or  of  abnormal  ty])es  in  variable  percentages.     Sperms 


47 

with  sluggish  motility  are  always  low  in  vitality,  and  have  weak 
powers  of  insemination,  as  the  motion  lasts  but  a  comparatively 
short  time.  The  vitality  may  be  but  moderately  lowered,  so  that 
although  the  sperms  are  highly  motile  when  ejaculated  they  soon 
lose  their  power  of  propulsion.  The  type  of  motion  is  likewise  an 
indicator  of  lack  of  vitality.  The  '"progressive  vibratilc"  motion 
described  by  Reynolds  should  proceed  to  a  high  degree  for  a  long 
period  before  the  "undulatory  tactile"  or  bunting  types  of  motion 
appear.  Early  appearance  of  these  two  latter  ty])es  indicates  in 
most  eases  a  marked  lack  of  vitality  of  the  elements.  The  motion 
should  be  vigorous  and  lasting,  for,  as  stated  by  Reynolds,  "noth- 
ing is  more  certain  than  that  spermatozjoa  of  merely  moderate 
vitality  seldom  impregnate  a  female." 

The  early  precipitation  out  of  the  "Boettcherchen"  crystals  is 
very  characteristic  of  oligospermia,  and  impotent  semen.  The 
theory  here  is  that  crystals  do  not  precipitate  out  wdien  a  fluid  i-^ 
actively  moving,  but  soon  do  so  when  the  fluid  is  motionless.  The 
sediment  which  normally  makes  up  about  two-thirds  of  the  sample 
is  usually  decreased  in  abnormal  samples.  The  semen  itself  should 
be  observed  for  unusual  viscosity  or  a  thin  watery  condition. 
Clotting  should  occur  readily  after  emission,  but  the  clot  soon 
liquefies  to  some  extent,  allow'ing  the  spermatozoa  to  become  more 
active.  This  clotting  is,  of  course,  to  protect  the  delicate  spei-ms 
from  the  hostile  secretions  of  the  vagina. 

The  presence  of  immature  and  deformed  types  of  sperms  rep- 
resents some  disturbance  of  spermatogenesis,  but  it  is  difficvilt  to 
explain  the  significance  of  these  forms.  They  are  seldom  seen  in 
normal  samples,  and  undoubtedly  none  are  capable  of  producing 
impregnation.  In  the  case  of  minor  abnormalities  of  staining  rf- 
actions.  the  sperms  are  probably  deficient  in  nuclear  material  or 
othei-wise  altered  so  that  probably  they  are  incapable  of  reaching 
and  uniting  with  the  ovum.  When  impregnation  does  occur  in 
the.se  cases,  weak  offspring  undoubtedly  result  in  many  instances. 

The  work  has  by  no  means  progressed  to  the  point  where  one 
may.  by  an  examination  of  the  semen,  determine  the  degree  of 
impotency  with  great  accuracy,  or  even  whether  the  animal  may 
be  restored  to  sexual  health  by  proper  hygienic  and  therapeutic 
treatment.  Ivelatively,  the  greater  the  changes  in  the  semen  and 
spermatozoa,  the  less  the  chances  of  impregnation.  Infertility 
to  any  marked  degree,  is,  however,  usually  accompanied  by  cor- 
responding ehanges  in  the  seminal  fluid  and  its  germinal  elements. 

Examination  of  the  semen  is,  and  pro])ably  always  will  be. 
simply  an  aid  in  reaching  a  diagno.sis.  AVhile  abnormalities  of 
the  semen  and  spermatozoa  are  associated  Avith  sterility  or  in- 
fertility, it  is  unwise  to  lay  too  much  emphasis  upon  this  method 
of  diagnosis  alone,  especially  Avith  regard  to  the  making  of  a 
definite  prognosis.  When  large  numbers  of  abnormal  spermatozoa 
are  present  in  the  semen,  we  are  safe  in  saying  that  the  animaJ 


48 

is,  at  the  time,  of  lowered  degree  of  fertility.  One  should  be  very 
cautious,  however,  in  foretelling  how  long  the  condition  will  last, 
or  if  the  animal  may  in  time  be  restored  to  full  fertility.  Ster- 
ility, due  to'  organic  disturbances,  probably  seldom  yields  to  treat- 
ment, but  when  it  is  due  to  functional  disorders  resulting  from 
defective  diet  or  lack  of  exercise,  the  condition  is  frequently  reme- 
died by  overcoming  the  cause.  Lack  of  exercise  and  overfeeding 
seem  to  be  etiological  factors  in  a  fair  percentage  of  cases. 

Besides  abnormalities  of  the  male  genital  tract,  we  must  always 
consider  the  numerous  factors  in  the  female  that  may  kill  or 
weaken  the  sperms.  Impediments  to  successful  coitus  may  be 
present  in  the  form  of  vaginal  constrictions,  abnonnally  short  or 
small  vagina,  or  other  deformities.  Hostile  exudates,  mechanical 
obstructions,  and  other  factors  may  interrupt  the  progress  of  the 
sperms  at  any  point  in  the  tract. 

Although  little  is  known  definitely  regarding  disorders  of  the 
endocrines  in  the  bull  and  their  relation  to  reproduction,  the  w^ork 
in  human  medicine  and  experimental  researches  upon  laboratory 
animals  warrant  thoughtful  consideration  of  these  factors  which 
are  by  no  means  insignificant.  In  the  future,  these  glands  will  no 
doubt  receive  more  and  more  attention  in  their  relation  to  the 
genital  organs  and  reproduction. 

Conclusions 

1.  The  genital  organs  of  the  bull  quite  frequently  undergo 
pathological  changes,  due  to  infection  with  the  same  varieties  of 
microorganisms  associated  with  genital  infections  in  the  female. 

2.  In  all  probability,  these  microorganisms  are  frequently  elimi- 
nated with  the  semen  and  infect  the  female  during  copulation. 

3.  Past  or  present  infection  in  the  genital  organs  of  all  the 
bulls  so  far  examined  was  evidenced  by  the  presence  of  the  fine 
connective  tissue  tufts  and  strands  upon  the  tunica  vaginalis,  par- 
ticularly that  part  covering  the  tail  of  the  epididymis. 

4.  LoAvered  sexual  capacity  is,  as  a  rule,  accompanied  by 
demonstrable  changes  in  the  semen. 

5.  A  study  of  impotency  and  sterility  includes  not  only  a  thor- 
ough study  "of  the  genital  organs,  but  also  those  extrinsic  factors 
which  govern  reproduction  either  directly  or  indirectly. 

6.  A  thorough  knowledge  of  the  anatomy  and  physiology  of  the 
male  genital  organs  is  fundamental  to  a  clear  understanding  of 
the  problem. 

I  am  much  imlebtecl  to  Drs.  W.  L.  and  W.  W.  Williams  for  some  of  the 
material,  and  for  helpful  co-operation  in  the  early  part  of  the  work;  to 
Drs.  C.  M.  Carpenter  and  R.  R.  Birch  for  many  helpful  suggestions;  and  to 
Dr.  J.  N.  Frost  and  others  who  so  kindly  co-operated  by  placing  at  my  dis- 
posal samples  of  semen  and  some  of  the  genital  tracts. 


49 


Bacteriology  of  the  Genital  Tracts  of  Normal  Young  Cal\^s 

Group  I 


Number 

Right  sem.  vesicle      Left  sem.  vesicle 

1 

Right  testis 

Left  testia 

1 

Staph,  albus. 

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p 

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Genital  Tracts  op 
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1 
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Description  of  Plates 
Plate  I. 

Fig.     1.     Diagrammatic  sketches  showing  the  development  of  a  spermato- 
zoon from  a  spermatogonium. 

Fig.     2.     Diagrammatic  sketch  showing  the  minute   structure  of  a  sper- 
matozoon.    The  middle  piece  is  made  comparatively  thick  in 
order    to    bring    out    the    finer    structures.      Adapted    from 
EUenljerger. 
Plate  II. 

Fig.     3.     Testicle  of  bull,  showing  extensive  degeneration  and  necrosis. 

Plate  III. 

Fig.     4.     Inner  part  of  wall  of  ductus  deferens.     Normal,     x  230. 

Fig.     5.     Same,   Imt   showing   extensive   degeneration   and   exfoliation    of 
the  lining  membrane,     x  230. 
Pt-Atb  IV. 

Fig.     6.     Ductus  deferens,  showing  entire  exfoliation  of  the  lining  mem- 
brane.    The  lumen  is  filled  with  a  cellular  debris,     x  50. 


61 

Fig.     7.     Same,   showing  the  degeneration  of  the  lining  membrane,  and 
debris  in  lumen,     x  230. 
Plate  V. 

Fig.  S.  Seminal  vesicle  of  bull.  High  power  section  showing  the 
normal  structure  of  the  vesicular  cavities. 

Fig.  9.  Seminal  vesicle  of  bull.  Low  power.  The  meml>rane  lining  the 
cavities  is  degenerated  and  exfoliate<i.  The  cavities  are 
filled  with  cellular  debris,  and  exudates.  There  is  some 
increase  in  the  interstitial  tissue,  and  atrophy  of  some  of 
vesicular  cavities. 
PXATE  VI, 

Fig.  10.  Low  power  section  of  seminal  vesicle  of  bull.  The  condition  is 
about  the  same  as  in  Fig.  9  except  that  it  is  not  quite  as 
severe. 

Fig.   11.     Same  as  Fig.  10.     High  power. 

Plate  VII. 

Fig.   12.     Testicular  tubule  showing  normal  spermatogenesis,     x  230. 
Fig.   Il3.     Testicular  tubule  showing  no  evidence  of  mitosis,     x  230. 
Fig.  14.     Testicular  tubule.     The  spermat-ogenic  epithelium  is  beginning 

to  degenerate  and  become  cast  off  into  the  lumen,     x  230. 
Fig.   15.     Testicular    tubule,    showing    almost    total    exfoliation    of    the 

spermatogenic  epithelium,     x  230. 
Plate  VIII. 

Fig.  16.  Testicular  tubule.  The  seminal  epithelium  is  entirely  degen- 
erated. The  membrana  propria  is  markedly  thickened. 
X  230. 

Fig.  17.  Testicular  tubule.  The  tubule  is  undergoing  atrophy  and 
degeneration.  The  interstitial  connective  tissue  is  much 
increased  in  amount,     x  230. 

Fig.   18.     Same  as  Fig.  17,  except  that  it  is  in  the  more  advanced  stages. 

Fig.   19.     Testicular   tubule.      The   spermatogenic   epithelium   has   under- 
gone a  sort  of  hydropic  degeneration.     The  interstitial  con- 
nective tissue  has   become   much    increased   in  amount   and 
has  undergone  cellular  infiltration. 
Plate  IX. 

Fig.  20.     Xormal  structure  of  epididymis  tubule,     x  140. 

Fig.  21.  Epididymis  tubule  showing  exfoliation  of  the  lining  membrane, 
and  cellular  debris  in  the  lumen,     x  140. 

Fig.  22.  Atrophy  and  degeneration  of  epididymis  tubule.  The  inter- 
stitial connective  tissue  is  much  increased  in  amount,    x  140. 

Fig.  23.     Same,  but  in  more  advanced  stage,     x  140. 

Plate  X. 

Fig.  24.     About  the  same  as  Fig.  22.     x  140. 

Fig.  25.     ilarked  degeneration  of  epididymis  tubule.    There  is  a  cellular 
infiltration  of  the  interstitial  tissue,     x  60. 
Pr.ATE  XL 

Fig.  26.  Spermatozoon,  showing  constriction  at  middle  of  head.  The 
head  is  also  somewhat  contorted,     x  670. 

Fig.  27.     Spermatozoon.     Pear  shaped  head,     x  670. 

Fig.  28.  Spermatozoon.  The  head  is  quite  long,  and  pointed  at  its 
posterior  end.     x  670. 

Fig.  29.     Spermatozoon,   showing  a  constriction   at  middle   of   the  head. 
X  670. 
Plate  XII. 

Fig.  30.     ^licrocephalic  spermatozoon,     x  670. 

Fig.  31.     Spermatozoon.     The  head  is  small,  and  pear  shaped,     x  670. 

Fig.  32.     Macrocephalic    sperm.      The    middle    piece    is    nuuli    thickened. 
X  670. 

Fig.  33.     Tailless  spermatozoa,     x  670. 


-F/ate  / 


A/udcU.% 

CentrosoTneS 
Ce-ntrosl:3h.ere 


QonnecUma  ??ec< 
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Discarded.  Cutobia'^m. 

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Gnter/or  Vodu-le. 
Neck 

'Posterior  nodule 
SbiraL  filament 
CutobLa^rnic  bkeath. 

Q-m-iixiar  nodlxic. 
l7?oolucrLL7n. 


PLATE  II 


Fig.  3. 


PLATE  III 


Fig.  4. 


Fig.  5. 


PLATE  IV 


Fig.  6. 


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Fig.  7. 


PLATE  V 


Fig.  8. 


Fig.  9. 


PLATE  VI 


Fig.   10. 


Fig.   11. 


PLATE  VII 


Fijr.   li 


1-ig.    VA. 


Fig.   14. 


Via.  15. 


PLATE  VIII 


Fig.   16. 


Fiff.   17. 


Fig.   18. 


Fig.   19. 


PLATI-:    IX 


Via.  20. 


Fig.  21. 


V\ii.   2-2. 


Fitr.  2;?. 


PLATE  X 


Fig.  25. 


PLATE  XI 


Fijr.  -ilj. 


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Fig.  29. 


PLATE  XII 


Fig.  30. 


Fig.  31. 


Fig.  32. 


Fig.  33. 


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